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FRONTISPIECE.—Viridian Dacnis, Mielero Esmeraldino, Dacnis viguieri. 


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Seyi THM SoONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 150, Part 4 


THE BIRDS OF THE REPUBLIC 
i OF PANAMA 
vA 


Part 4.—Passeriformes: 
Hirundinidae (Swallows) to 


Fringillidae (Finches) 


By 
ALEXANDER WETMORE 


Research Associate (Deceased) 
Smithsonian Institution 


ROGER F. PASQUIER 


International Council for Bird Preservation 
Smithsonian Institution 


AND 


SLORKS E) OLSON 


Curator, Division of Birds 
Smithsonian Institution 


SAATHSON/Apy 


FEB 26 1985 


LIBRARIES 


SMITHSONIAN INSTITUTION PRESS 
City of Washington 
1984 


Copyright © 1984 by the Smithsonian Institution. All rights reserved. 
Published in the United States by the Smithsonian Institution Press. 


Library of Congress Cataloging in Publication Data 
(Revised for volume 4) 


Wetmore, Alexander, 1886— 
The birds of the Republic of Panama. 


(Smithsonian miscellaneous collections ; v. 150) 

Pt. 4 by Alexander Wetmore, Roger F. Pasquier, and 
Storrs L. Olson. 

Vols. 1-2: Publication 4617, 4732. 

Includes bibliographical references. 

Supt of Docs; no.: Sl 12=B53/27/pr4 

CONTENTS: 1. Tinamidae (tinamous) to Rynchopidae 
(skimmers).—pt. 2. Columbidae (pigeons) to Picidae 
(woodpeckers) .—[ etc. |—pt. 4. Order Passeriformes, 
suborder Passeres, Hirundinidae (swallows) to Fringillidae 
(finches). 


1. Birds—Panama—Collected works. I. Pasquier, 
Roger Fil) (Olsony Storrs ie i iitles Vee Senitess 
OlMS7 Vvolals0eete: 598.297287 66-61061 
ISBN 0-87474-122-X (v. 3) 


Part 1 of The Birds of the Republic of Panama by Dr. 
Alexander Wetmore was published by the Smithsonian 
Institution in 1965 as volume 150 of its Miscellaneous 
Collection series. It covers in systematic sequence the 
Tinamidae (Tinamous) through the Rynchopidae 
(Skimmers). 


Part 2 of The Birds of the Republic of Panama by Dr. 
Alexander Wetmore was published by the Smithsonian 
Institution in 1968 as volume 150, Part 2, of its Miscel- 
laneous Collections series. It covers in systematic sequence 
the Columbidae (Pigeons) through Picidae (Wood- 
peckers). 


Part 3 of The Birds of the Republic of Panama by Dr. 
Alexander Wetmore was published by the Smithsonian 
Institution in 1972 as volume 150, Part 3, of its Miscel- 
laneous Collections series. It covers in systematic sequence 
the Dendrocolaptidae (Woodcreepers) to Oxyruncidae 
(Sharpbills). 


Introduction 


CONTENTS 


aéu'aiie, (eke) e)ie 70) (eo) sal elie Lela), e@ .@) 610) oe: 76) (8) 88) (6/18). 6) \8 (0) 8) 18) ev ole! (8) 66 (a (0) 8)) ofa) e016) 16 Ke 


MMT Gl TTT E TIE Seep heer e ere rents Aes Sache ant ai ala iat Be hy wn) 5S Ugh og Sal 


EEPROM OR MeBSI a! phan c eo Sie asen tase oh ahem ay sabentelishae, Oe eae decane ae 


Family 
Family 
Family 
Family 
Family 
Family 
Family 


Family 
Family 


Family 
Family 
Family 
Family 


Family 
Family 


Family 


Family 
Family 


Liiruneimidae: swallows. Golondrinas 2257 Ge. <2. 4-6 
Corvidae: Crows and Jays, Cuervos y Urracas ....... 
Cmchdac- Wippers, Minlos, AcHaticos.)) 52260 a iieten ns ee 
iicoclodytidac-)VWinens, Cucarachetos, 2.7)... usc 
Mimniddes Mhrashers, Mockingbinds 22i2-224s24 205... 
sninaidae seb mnushes; Zorzahes ibiniice. SA ase seers wir 


Ptilogonatidae: Solitaires and Silky-flycatchers, 
Solitamosys Wordles “Sedososm-ics «sons wie eater neo ays 


Sylviidae: Gnatwrens and Gnatcatchers, Cazajejenes .. 


Motacillidae: Pipits and Wagtails, 
Bismitasmy mMbavandenrdSt ens isl. scr. Gale = lee eee sere de 


Bombyellidae: Waxwines, Chinitos .2-..2:2.--.-..- 
Mioceidde- \Veaver inches, minzones Mejedores...4-. - 
Strnidae:  Stamines. Wi stominos: 2). 22 sees cineca d 


Vireonidae: Vireos, Peppershrikes and Shrike-Vireos, 
NWAthe@ Seavae uOll cle hOStus tenance hie een ony ee. 8 


Barulidaec Wood: Warblers, Remitas, .25. 002... 5... - 


Icteridae: Blackbirds and Orioles, 
dhinpialesmy ACUACATCLOSe 6 58 SCN eka se hearse 


Thraupidae: Tanagers, Tangaros, 
MD AMASEASH Y MEMMUITCHOS eveio tara k ales ctaheyers sie cua ds 


Coerebidae: Honeycreepers, Mieleros ............... 


Fringillidae: Finches, Gorriones y Pinzones ......... 


RenpeMGina.) Nddenda ito (Volumes 13, a0. sea oe cs sks vale wis e's 


639 


Mist (Or TMicw St RA Tons 


FRONTISPIECE— Viridian Dacnis, Mielero Esmeraldino, 


Dacnis vigumert. 


FIGURE 


1 
z: 


oe 


Mangrove Swallow, Golondrina Manglera, Tachycineta albilinea 
White-thighed Swallow, Golondrina Muslos Blancos, Neochelidon 
NO BNES. CONTEC NE TATE Se SRS DE UMMA EAN GEV OTe en Le 
Portion of outer primary feather of the Rough-winged Swallow, 
Golondrina Ala de Sierra, Stelgidopteryx ruficollis, showing 
lrora anime ahi) arb Shays Mies bn SoNM RLM asa cae su. 4 


. Black-chested Jay, Chocho, Cyanocorax affims geledom ........ 
. Silvery-throated Jay, Urraca Garganta de Plata, Cyanolyca ar- 


IG NU ARGC TULUG ULC Ny ey EVN SAIN Ae gia tale Ma iy SMa au 8 oaliauee 


. American Dipper, Mirlo Acuatico Americano, Cinclus mexicanus 


LE GHLGIES ENC MUM On LNAI ODB A MTLE TV aGU i ay AG SMI SU ect 


. Black-capped Donacobius, Paralauta de Agua, Donacobius atri- 


COPS: HOCH DEE ISN ER USA Cann eae) EA ARR  Monie nia Ry tren re 


. White-headed Wren, Cucarachero Cabeciblanco, Campylorhyn- 


CPS: HOD TLE ISS MALATESTA DU COMO Ue Se NUE TAL RG aa NG aa 


. Band-backed Wren, Cucarachero Listado, Campylorhynchus 


MO AUGACOSEOMUGEIESTS Mi un NN Te oe LU RIC NY AAACN NEE ash tele 


. Bay Wren, Cucarachero Cabecinegro, Thryothorus nigricapillus 
. White-breasted Wood-wren, Cucarachero Pechiblanco, Hent- 


BOM CLUGO SEDC LEW nied Ss ick ese n egy Us bey hacia sie nen ape ean ne INI 8 | 


. Slaty-backed Nightingale-thrush, Zorzal Pizarrefio, Catharus 


[PUSCORCE® SINAN UE ECG ISIS AIR OIE SE HARA SDN OUR ONAN 


. Black-faced Solitaire, Solitario Carinegro, Myadestes melanops 
. Long-tailed Silky-flycatcher, Capulinero Colilargo, Ptilogonys 


COMME SS AMAIA NIN CONES AVN VAIS EE cE cu TAO AUT aN Rats 


. Tropical Gnatcatcher, Cazajején Tropical, Polioptila plumbea, 


eR ADElOW) r teltaley CADOVE)) seis core ec len nie ave a Ura L. 


. Yellowish Pipit, Bisbita Amarillenta, Anthus lutescens parvus. 
. Cedar Waxwing, Chinito Cedroso, Bombycilla cedrorum ....... 
. Rufous-browed Peppershrike, Pajaro Perico, Cyclarhis gu- 


A DIEOLA STS! BN SUITS SERIE AIG SO EO DS Mn PNR IN gS i AA OR 


. Green Shrike-Vireo, Follajero Verde, Smaragdolanius pulchellus. 
. Yellow-winged Vireo, Vireo Aliamarillo, Vireo carmioli ....... 
. Scrub Greenlet, Verdecillo Rastrojero, Hylophilus flavipes. .... 
. Black-and-white Warbler, Reinita Trepedora, Mniotilta varia, 


IAMS ee Ae SMa UR SR UNCC nO GUN Yt Mg A SEO CUTE MRM RR 


110 


154 
168 


a7 


179 
191 
195 


199 
205 
ZN2 
238 


V1 


Zoe 


BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Yellow Warbler, Reinita Mangletera, Dendroica petechia, a male 
of the erithachorides (Mangrove Warbler) group. -..-.2.22. 


. Collared Redstart, Candelita Collareja, Myioborus torquatus ... 
. Buff-rumped Warbler, Reinita de Rabadilla Anteada, Basileuterus 


fulviCaUda. oe. eh RR a ee ee 


. Zeledonia, Zeledonia, Zeledonia coronata ......... 25. 
. Montezuma Oropendola, Chacarero de Montezuma, Psarocolius 


WMONTECZUING oa es hacen Bin 8 ORR 5 eas ake en See 


. Chestnut-headed Oropendola, Chacarero Comtun, Psarocolius 


waglert; male 6.0.6.2. he AE, See 


. Giant Cowbird, Vaquero Gigante, Scaphidura oryzivora, male .. 
. Bronzed Cowbird, Vaquero Bronceado, Molothrus aeneus aeneus, 


female Ceft),’male (right)! 2s.) ae oe 


. Yellow-tailed Oriole, Parao Coliamarillo, Icterus mesomelas ... 
. Swallow-Tanager, Azulejo Golondrina, Tersina viridis occiden-— 


tals, male te ao ee. oe ne 


. Spot-crowned Euphonia, Tanagra Coronimanchada, Euphonia 


imioans, male (above), temale (below) 2)... 47 eee 


. Spangle-cheeked Tanager, Tangaro Carirrayado, Tangara dow 
. Crimson-backed Tanager, Sangretoro Comtn, Ramphocelus 


dimidiaius, male (above); female (below) ©. =.. 2.5 .cmeee 


. Rosy Thrush-Tanager, Frutero Rosaceo, Rhodinocichla rosea exi- 


MAD eS ORE ETE UES 7 


. Black-and-yellow Tanager, Frutero Orinegro, Chrysothlypis chry- 


somelas, male” (below), female (above)... 2... 75s eeeeee 


; Bananaquit, Mielero Platanero, Coereba jlavcola. . 7. ae 
. Slaty Flower-piercer, Mielero Picaflor, Diglossa baritula plumbea, 


male. ee oO EE 2G AE 


. Scarlet-thighed Dacnis, Mielero Patirrojo, Dacnis venusta, male 
. Black-headed Saltator, Saltator Cabecinegro, Saltator atriceps 


lacertosus oo. oe ee ee ee eee 


. Buff-throated Saltator, Saltador Gargantianteado, Saltator ma-i- 


Mus 


© 0 © © © e@ 0 8 © « ¢ © 6 (0 0 «© ©. (6 © 6 0 0 © © © « 0 0 © 0 « «, © ©) 6 © 6 © ‘e je fe 6) (s) (6) [s) eee) salle 


. Streaked Saltator, Saltador Pechirayado, Saltator albicollis .... 
. Slate-colored Grosbeak, Piquigrueso Apizarrado, Pitylus grossus 


Saturatus, Male oo) oss onde wl ees ee eee 


. Blue-black Grassquit, Arrocero Piquiagudo, Geospiza jacarina 


splendens, temale (lett); males (cight). 20/2 22) 


. Distribution of the races of Sporophila americana in Panama 


(from Olson, 1981)" 27) Ss, oS ee 


. Volcano Junco, Junco del Volcan, Junco vulcamt ............-. 
. Rufous-collared Sparrow, Gorrién de Collar Rojizo, Zonotrichia 


capensis’ Costaricensis oo POR ee eee 


. Wedge-tailed Grass-finch, Pinzén Yerberero Coliacufiado, Em- 


berizoides herbicola hypochondriacus .....2.00n00+00++08008 


260 
312 


326 
Sl 


337 


342 
356 


399 
370 


389 


406 
432 


441 


484 


488 
501 


506 
a3 


ao i 


533 
536 


545 


558 


567 
610 


612 


THE BIRDS OF THE REPUBLIC OF PANAMA 
PART 4: ORDER PASSERIFORMES, SUBORDER 
PASSERES, HIRUNDINIDAE (SWALLOWS) 
TO FRINGILLIDAE (FINCHES) 


By ALEXANDER WETMORE 
Research Associate, Smithsonian Institution (Deceased) 


Rocer F. PASQUIER 
International Council for Bird Preservation, 
Smithsonian Institution 
and 
Storrs L. OLson 
Curator, Division of Birds, Smithsonian Institution 


INTRODUCTION 


As with so many ambitious works in ornithology, Alexander Wet- 
more’s Birds of the Republic of Panama unfortunately required more 
than a lifetime to complete. Those of us around the Division of Birds 
who were privileged to work with Dr. Wetmore could only hope that he 
would live to see his magnum opus finished, but even his indefatigable 
constitution inevitably gave way to the attrition of time. He was 86 
years old in 1972, when Part 3 appeared, and it is a tribute to his en- 
durance that he proceeded as far with the manuscript for Part 4 as he 
did. When his health failed, he had prepared accounts for almost all 
of the “ten-primaried oscines.” Alexander Wetmore died at his home 
on December 7, 1978, at the age of 92 after a long illness. 

Dr. Wetmore’s colleague and friend of long standing, S. Dillon Rip- 
ley, Secretary of the Smithsonian Institution, approached me, when it 
became certain that Dr. Wetmore would be unable to continue his 
studies, to inquire into the feasibility of completing Part 4 of Birds of 
the Republic of Panama. I agreed to be responsible for the identifica- 
tion of specimens and for systematic decisions, provided someone else 
could be found to compile the species accounts and descriptions that 
remained to be done. Happily for all concerned, we were able to enlist 
the services of Mr. Roger F. Pasquier, who has attended this task with 
determination and assiduity. 

In addition to the large collections that Dr. Wetmore himself had 
made in Panama, he worked simultaneously upon extensive collections 


I 


2 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


from Colombia obtained by M. A. Carriker. When I began work on 
the project, there still remained thousands of specimens of nine- 
primaried oscines and a few other families from Panama and Colom- 
bia that had not been identified or incorporated in the main collections. 
I have evaluated the geographic variation in each of these remaining 
species and have made all of the decisions about subspecific limits. In 
some instances I was not able to recognize taxa that are currently held 
to be valid, whereas in others I was able to discern undescribed sub- 
species. Most major innovations I have proposed elsewhere in technical 
journals. I have made no attempt to try to guess how Dr. Wetmore 
might have interpreted each species; therefore, the systematics used 
in the section from the Ptilogonatidae onward is mine alone. An ex- 
ception is made for the subspecies of North American migrants, where 
I have deferred to the judgment of my colleagues John W. Aldrich, 
M. Ralph Browning, and Allan R. Phillips, each of whom has been 
consulted regarding particular taxa. 

Any sequence of species is bound to be to some extent arbitrary. 
Because Ridgely (1976) followed the sequence that Dr. Wetmore had 
used in the first three volumes of this series, we have simply adopted 
the sequence of species in Ridgely for the present volume, in order to 
make the two works compatible. Exceptions are Donacobius, now 
placed at the beginning of the Troglodytidae, Myadestes, which ap- 
pears in the Ptilogonatidae, and Spiza, which now leads off the Icter- 
idae. The artificial family ‘“Coerebidae”’ we have placed after the 
Thraupidae, in which family most of the coerebid genera properly be- 
long. Although we have included the Cyclarhidae and Vireolaniidae in 
the Vireonidae, and the Zeledoniidae in the Parulidae, this has not af- 
fected the sequence of species. 

As mentioned, Dr. Wetmore had completed his studies on the taxa 
up to the Ptilogonatidae, and this portion of the text we have attempted 
to leave as intact as possible, although the effort has been made to in- 
clude information published since the manuscript was originally com- 
pleted. Mr. Pasquier has assembled the species accounts from the field 
notes and bibliographic index left by Dr. Wetmore and from the sub- 
sequent literature. Because so much of the information comes directly 
from Dr. Wetmore’s field notes, the first person singular has been re- 
tained throughout, which also conforms with the first part of the vol- 
ume, written by Dr. Wetmore alone. In places I have interjected my 
own comments, these being initialled and set off in brackets. Citations 
of “in ltt.” refer to correspondence received by either Pasquier or Ol- 
son, unless indicated otherwise. From the Ptilogonatidae onward, all 


INTRODUCTION 3 


descriptions and measurements were made by Mr. Pasquier. Spanish 
names through the Ptilogonatidae are from Dr. Wetmore’s manuscript, 
with occasional modification by Eugene Eisenmann; the remaining 
Spanish names were kindly provided by Drs. Pedro Galindo and Eu- 
storgio Mendez of the Gorgas Memorial Laboratory in Panama. A\I- 
though the Canal Zone no longer exists as a political entity, we have 
continued to use this designation to reflect the localities that appear on 
specimen labels and to be consistent with the remainder of this series. 

Several species belonging to families treated in the first three vol- 
umes have subsequently been found in Panama for the first time. Ac- 
counts of these species appear in an appendix. As in the earlier volumes, 
accounts of species whose presence in Panama has not been supported 
by specimens are bracketed, and literature citations appear within the 
text. We have resorted so frequently to Ridgely’s indispensible Guide 
to the Birds of Panama (Princeton University Press, Princeton, N. J., 
1976), that this reference is cited simply as “Ridgely, 1976,” with the 
appropriate page number. 

Dr. Wetmore originally intended to include a complete bibliography 
as well as an account of the history of ornithology in Panama, but we 
have not attempted to provide either and have settled simply for finish- 
ing the species accounts. A gazetteer is in preparation to be published 
separately. As we could not hope to emulate the devotion that Dr. Wet- 
more invested in the production of the first three volumes in this series, 
we can only trust that the ultimate result will be judged better than 
nothing at all. 


ACKNOWLEDGMENTS 


We are especially indebted to the late Eugene Eisenmann and to 
Robert S. Ridgely, who read over drafts of the manuscript and used 
their special knowledge of Panamanian birds to suggest many valuable 
alterations and additions. We have also had the pleasure of consulting 
frequently with Mrs. Alexander Wetmore concerning various aspects 
of the production of this volume. Further information and specimens 
that were of use in the preparation of this volume were provided by John 
Farrand, Jr., John W. Fitzpatrick, Pedro Galindo, Frank B. Gill, John 
W. Hardy, C. Lynn Hayward, Ned K. Johnson, Lloyd F. Kiff, Wesley 
I. Lanyon, Marcy Lawton, Eustorgio Mendez, John P. O’Neill, Ken- 
meth) ©: Parkes, Allan 'R.) Phillips, Ralph W. Schreiber, Robert W. 
Storer, and Melvin A. Traylor, to all of whom we remain indebted. 

Although many of the text drawings for this volume had already 


4 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


been executed by the late Walter Weber, certain prominent taxa were 
lacking. We were fortunate to have Guy Tudor to remedy the defi- 
ciencies and also to render the color frontispiece. Finally, it should be 
recognized that were it not for the vigorous support and warm en- 
couragement of S. Dillon Ripley, this volume would never have been 
completed. 

SToRRS L. OLSon 

Smithsonian Institution 

Washington, D.C. 

April 1982 


Order PASSERIFORMES 
Family HIRUNDINIDAE: Swallows, Golondrinas 


The swallow family, almost worldwide in its distribution, has 11 
species found regularly in Panama. All are birds of active flight that 
feed on insects captured mainly on the wing. Winter migrants from the 
north include 6 kinds, one of them, the Rough-winged Swallow, also 
represented by a local race that nests widely through the Isthmus. Two 
others, less numerous, are recorded as visitors from South America 
during the months of southern winter, one of them—the Blue-and- 
white Swallow—also with a resident subspecies in the western high- 
lands. The Southern Martin of southern South America is definitely 
known from a single specimen record in eastern San Blas. The Cave 
Swallow is reported only from a specimen of uncertain history. 

Migrant flocks of swallows during the months of northern winter 
may include large numbers of a single kind, but it is usual to see 2 or 
more species in mixed company, joined where insects are abundant 
through this common source of food. While most common in the low- 
lands, swallows may be found in open lands at any elevation. The north- 
ern migrants, especially the Barn Swallow, most abundant in western 
Panama, often gather at dusk in large groups to sleep in growths of 
rushes in marshy areas or in sugarcane. Mixed with others of the 
family, they range in companies through the day. The Brown-chested 
Martin, a southern season migrant, late April to October, from Brazil 
and Argentina, gathers at night to sleep in trees (at least in its breeding 
range) in large assemblages. It appears that this species, now common, 
may have extended its winter range to Panama within recent years, 
as it was not reported by the early naturalists who made collections and 
observations in its present isthmian range. 


10. 


1 


EZ. 


FAMILY HIRUNDINIDAE 5 


KEY TO SPECIES OF HIRUNDINIDAE 


BeESELGONVi MO: trace Of DIME. fan. secu oe coe wenn oe ce ee ee anes 2 


DHChesteel Oley Or reel att teehee eo ee ee ee) ke 5 
Undersurface entirely sooty brown, except thighs. 
White-thighed Swallow, Neochelidon tibialis minima. p. 29 
Pied USA Gey Pan tiellliy | WiMlbes .pshorae yo eusetahew i orcs trteeas Pret ianeke Were b kek 3 
Outer edge of outermost primary serrated; no distinct breast band. 
Rough-winged Swallow, Stelgidopteryx ruficollis. p. 36 
Outer edge of outermost primary entire; distinct dark breast band...... 4 
Throat white, total length 125 mm or less. 
Bank Swallow, Riparia riparia riparia. p. 5 
Throat white, total length 155 mm or more. 
Brown-chested Martin, Progne tapera fusca. p. 27 
Back bright metallic green. 
Violet-green Swallow, Tachycineta thalassina lepida. p. 11 
Poe mcubinely Onmipantly steel blues 20. 02h ee wee a es 6 
Undersurface cinnamon-rufous. 
Barn Swallow, Hirundo rustica erythrogaster. p. 13 
Piidenstnnace wilike vor Steel) DIG: sc. side c oes shoe ee eee ee eas bee 7 
Rump buff to brown. 


ama MS KeCMMDIIe: OF CWither 20.40 so00 GAR ee be RS BOE et RR 8 
Rump white. 


SS AISEND CIBER AS SRINESE TS SIRs a PF ORE i gen ate ee Ua 9 
ET AERCORAE TULUM a Pay GRR ae ee ee he on 10 
Mie abestcel bite or wath: brows oes. Shh Dae ee a 11 


Larger, total length 130-140 mm. 
Tree Swallow, Tachycineta bicolor. p. 7 
Smaller, total length 110-125 mm. 
Blue-and-white Swallow, Notiochehdon cyanoleuca. p. 32 
Undersurface steel blue. 
Purple Martin, Progene subis,* male. p. 22 
Minderstmrace DROMMrana WiIhten less e leva Close oe ees Be Pas ie Fe 12 
Forehead feathers tipped gray or whitish. 
Purple Martin, Progne subis,* female. p. 22 
Forehead steel blue or brown. 
Gray-breasted Martin, Progne chalybea chalybea. p. 19 


RIPARIA RIPARIA RIPARIA (Linnaeus): Bank Swallow, 
Golondrina Barranquera 


Hirundo riparia Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 192. (Sweden.) 


Small; undersurface white, with a distinct grayish brown band across 


the lower foreneck and upper breast. 


*There is also one specimen record of the Southern Martin, Progne elegans, 


from Panama. For comparison with P. subis see text, p. 25. 


6 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Description —Length 115-125 mm. Adult (sexes alike), upper sur- 
face of body, including side of head, and tail grayish brown; forehead 
slightly paler than crown; back also usually somewhat paler; scapulars, 
tertials, rump, and upper tail coverts with lightly marked paler margins; 
wings, including coverts, darker (blacker); undersurface, including 
sides and undertail coverts, white; a broad band of a grayish brown 
across the foreneck and upper breast, continued laterally on the upper 
sides. 

Immature, rump, upper tail coverts, tertials and wing coverts tipped 
with cinnamon-buff; chest band also with paler tips. 

Measurements.—Males (10 from New Jersey, Rhode Island, Iowa, 
Nevada, and Alaska), wing 99.7-105.7 (102.0), tail 47.0-51.7 (49.1) 
culmen from base 8.0-8.9 (8.3), tarsus 10.0-11.1 (10.4) mm. 

Females (10 from New Jersey, District of Columbia, Louisiana, 
Kentucky, Illinois, Idaho, and Ontario), wing 94.4-102.1 (98.0), tail 
42.6-49.6 (47.6), culmen from base 7.0-8.7 (7.8), tarsus 9.9-11.7 
(10.6) mm. 

Winter visitor from the north, September to April, found irregu- 
larly, often with flocks of other swallows. The first fall migrants may 
appear early, as Ridgely (1 litt.) reported 2 at El Llano, eastern Prov- 
ince of Panama on August 2, 1974, Arbib and Loetscher recorded 1 
at Gatun, Canal Zone, August 26, 1934, and Eisenmann reported some 
seen with other swallows on the savannas of Coclé, September 1, 1954. 
There are numerous records through September and October, when 
they appear still to be in southward passage. Specimens in the British 
Museum were taken by H. J. Kelsall at sea in the Gulf of Panama on 
October 4 and 25, 1924. Griscom recorded 3 taken October 2, 6, and 17, 
1930, at Puerto Obaldia on the eastern San Blas coast. 

A few may be seen through the winter period, as Eisenmann recorded 
several at Playa Coronado, November 25, 1962, and Jewel saw them 
in the Canal Zone on November 30, 1911, and February 22, 1912. The 
latter date may mark the beginning of the migration northward, during 
which lesser numbers are recorded in March and April. Late dates of 
small numbers, presumably stragglers, are reported for May 11, 15, 22, 
27 (Ambrose, Ridgely, N. G. Smith) and June 3, 1961 (Gatun Dam, 
Ambrose, 10 birds). On March 14, 1944, I noted several over the Gulf 
of Panama, in crossing from Isla San José to Balboa. Griscom re- 
corded 10 on March 8, 1927, a mile offshore from Cabo Garachiné. 
Two males in the Hancock collections of the Los Angeles County Mu- 
seum were collected April 27, 1939, from the deck of the yacht Velero 
III in Bahia Caledonia, on the eastern San Blas coast. 


FAMILY HIRUNDINIDAE oh 


Usually Bank Swallows are noted in company with other migrants 
of the family, less often alone in groups of up to 100 individuals. Dur- 
ing the winter of 1975-76, a year of exceptional swallow flights in 
Panama, Ridgely and J. J. Pujals noted about 25 on January 27 at 
Tocumen, eastern Province of Panama, with thousands of Barn Swal- 
lows that were definitely not migrating. P. Donahue saw 2 on January 
2, 1980, at Aguadulce, Coclé (Eisenmann in /itt.). 


TACHYCINETA BICOLOR (Vieillot): Tree Swallow; Golondrina Bicolor 


Hirundo bicolor Vieillot, Hist. Nat. Ois. Amer. Sept., vol. 1 (1807), 1808, p. 61, 
pl. 31. (New York.) 


Medium size; undersurface white; adult, above steel blue; immature, 
duller, browner above; differs from the Mangrove Swallow in lack of 
white on upper surface, and larger size. 

Description.—Length 130-140 mm. Male, upper surface, including 
lesser wing coverts, greenish steel blue; middle wing coverts black, 
edged with greenish to steel blue; rest of wings and tail dusky; lores 
black; undersurface white; axillars and underwing coverts brownish 
gray; edge of wing white. 

Female, similar but usually duller colored above; breast in some 
washed with brownish gray. 

Immature, upper surface dark mouse gray; usually with breast 
washed with grayish brown; tertials, and in some individuals, outer 
upper tail coverts, edged narrowly with white to grayish white. 

Measurements.—Males (10 from Florida, winter), wing 115.0-125.2 
(118.4), tail 52.1-58.0 (54.7), culmen from base 9.0-11.2 (10.2), tarsus 
12.0-14.4 (13.1) mm. 

Females (10 from New York, Florida, winter, spring), wing 108.8- 
117.0 (112.2), tail 47.8-53.6 (51.5), culmen from base 8.2-10.9 (9.2), 
tarsus 11.1-13.2 (12.3) mm. 

Winter migrant from the north; irregular, recorded chiefly in the 
Caribbean coastal lowlands of western Bocas del Toro, and in the Canal 
Zone. 

The Tree Swallow, an abundant species over a wide range in the 
north, is common during the winter months south to eastern Honduras. 
Farther south, along the eastern coast of Central America, it is little 
known. 

On January 17, 1958, more than 100 Tree Swallows ranged with 
other swallows over open lands partly flooded from recent rains near 
Changuinola, Bocas del Toro. I collected a male there on March 4 


8 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


from several flying with Barn and Rough-winged Swallows. In the 
Canal Zone, Eisenmann on February 1, 1973, found a small flock near 
Gamboa, and on February 4, a larger group of 200-300 at the Gatun 
Dam spillway. On February 16, 1969, Ridgely (1976, p. 259) photo- 
graphed Tree Swallows that were part of a flock of 225 at the Gatun 
Dam. In the early months of 1975 and of 1976 flocks were seen in vari- 
ous locations in the Canal Zone; Ridgely (1m litt.) found as many as 
600 at La Jagua on January 27, 1976. The latest date for which the 
species is recorded in Panama is April 5, when in 1975 Eisenmann (in 
htt.) and others saw 6 (2 adults) at Gatun Lake. 

Tree Swallows are not found in Panama every year, but the species 
has recently been seen in northern Colombia (20 at Salamanca Na- 
tional Park, Magdalena, February 6, 1977, by Ridgely) and in northern 
Venezuela (at Chichiriviche Lagoon, Falcon, by P. Alden in 1972 and 
1976), suggesting that they often travel farther south than was pre- 
viously known. 


TACHYCINETA ALBILINEA (Lawrence): Mangrove Swallow, 
Golondrina Manglera 


FIGuRE 1 


Petrochelidon albilinea Lawrence, Ann. Lyc. Nat. Hist. New York, vol. 8, May 
1863, p. 2. (Atlantic slope, near the Panama Railroad, Canal Zone, Panama.) 


Small; undersurface, rump, line above lores, and tips of secondaries 
white. 

Description.—Length 105-115 mm. Adult (sexes alike), crown, 
hindneck, back, scapulars, lesser wing coverts, and upper tail coverts 
steel blue, with a greenish to bluish sheen (greener when in fresh plum- 
age); a narrow line above lores white; lores and narrow line beneath 
the eye black; wings and tail dull black; tertials and middle wing coverts 
edged narrowly with white; lower hindneck with feathers white basally, 
this reduced in some to an indistinct white line; rump white, with 
finely marked dusky shaft lines; lower surface, including underwing 
coverts, white, with the breast and sides faintly washed with pale gray. 

Immature, lores dull black, bordered above by a narrow white line; 
rump white; rest of upper surface, with the wings, drab gray; under- 
surface white, with a faint wash of somewhat brownish gray on breast 
and sides; outer underwing coverts banded indistinctly with dull gray. 

A female collected at Canita, on the Rio Bayano above El Llano, 
February 6, 1962, had the iris wood brown; tarsus and toes fuscous- 
black; claws black. In another female, at Puerto Armuelles, Chiriqui, 


FAMILY HIRUNDINIDAE 9 


February 14, 1966, the iris was dark brown, bill black, tarsus and toes 
fuscous-brown; claws black. A third, from the second locality, March 
9, 1966, had the tarsus, toes, and claws black. In this third bird the 
tongue was flesh color, and the lining inside the mouth was black. 

Measurements.—Males (10 from Panama), wing 93.4-101.1 (96.8), 
tail 39.6-42.2 (40.8), culmen from base 9.9-11.8 (11.2), tarsus 9.5- 
15 -C11.2) mm. 

Females (10 from Panama), wing 92.2-98.2 (94.7), tail 39.8-46.8 
(42.0), culmen from base 10.0-11.6 (10.8), tarsus 10.1-11.5 (10.7) 


mm. 


FicurE 1.—Mangrove Swallow, Golondrina Manglera, Tachycineta albilinea. 


Resident. Common throughout the lowlands, mainly along the 
larger rivers and over open water around the coastal mangrove swamps; 
seen regularly along the Panama Canal, on Gattin Lake, and in bays on 
the coasts, usually near the mouths of the rivers. Rare in the highlands. 

Recorded on Islas Verde and Cébaco in Golfo de Montijo; Isla Coiba. 

From January to May these small swallows are seen in pairs or little 
groups; in the second half of the year they sometimes gather in larger 
groups of 50 or more. They range over open water along the larger 
streams, often resting on fallen branches or logs stranded in the water. 
A few may be seen flying along beaches or over pastures and other 


IO BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


open lands, but they never seem to stray far from water. They seem to 
wander rarely to the upland country. My only record there is of one 
seen at a small lake near El Volcan, March 20, 1960. 

The nesting season appears to extend from January to June. A pair 
taken over the Rio Chagres at Juan Mina, Canal Zone, December 14, 
1955, while ranging attentively together, showed no development of the 
sexual organs. Others in January at several localities were breeding 
individuals. Grown young, recently from the nest, are common in June. 
They nest in holes in snags and stubs, fallen or standing in the water, 
sometimes in cavities made originally by woodpeckers. The usual lo- 
cation is low but above flood level. In Gattin Lake they may use the 
metal cans placed at the tops of channel markers. 

January 31, 1957, at Mandinga, San Blas, I found a nest placed in a 
shallow cavity in the end of one of the pilings supporting an ancient 
wharf in the sheltered bay, when the female swallow flew out as we 
passed near in a dugout canoe. The bulky mass of grass and other 
plant stems that filled the lower part of the cavity held a deep cup of 
finer materials, lined with the white and gray contour feathers of 
herons. The 4 eggs, about one-third incubated, were oval in form and 
white in color, without markings. They measure 18.7 x 12.6, 19.0 12.6, 
19.2*12.7, and 19.4X12.5 mm. \Aset of 2 in the British iiiseam 
(Natural History) sent to Salvin and Godman, by F. Blancaneaux 
from British Honduras, collected May 5, 1888, also are oval, and white 
without gloss. They are slightly smaller, as they measure 17.7X13.1 
and 18.0X13.4 mm. Russell (A. O. U. Mon. no. 1, 1964, p. 132) says 
that Peck, in British Honduras, in addition to the usual cavities in 
stumps standing in water, also found nests placed “in abandoned wood- 
pecker holes in pine trees situated in very open areas of the lowland 
pine ridges.” 

Stomachs of a few examined were filled with remains of a variety of 
small insects, including diptera, hymenoptera, coleoptera, and hemip- 
tera (see also Ricklefs, Auk, 1971, pp. 635-657). A female collected by 
Strauch (Bull. Brit. Orn. Club, 1977, p. 64) weighed 6.4 g. 

At Juan Mina, in January, in evening when Barn and other swallows 
were moving over the river to communal roosts in marshy areas, I 
found 20 or so of the present species gathered by themselves to sleep 
on a many-branched dead snag standing isolated in the river. 

The species is one of extensive range from southern Sonora and 
southern Tamaulipas, Mexico, south along the coastal lowlands through 
Central America to eastern Panama, where it is recorded east to the 
mouth of the Rio Mandinga on Golfo de San Blas, western San Blas, 


FAMILY HIRUNDINIDAE Let 


on the Atlantic slope; to Boca de Cupe, above El Real on the Rio Tuira, 
and above the mouth of the Rio Tuquesa, on the Rio Chucunaque, on 
the Pacific side. In July 1975, Ridgely (1 litt.) found this species quite 
common along the lower Chucunaque ( Yaviza area) and Tuira Rivers, 
but it became relatively scarcer as he proceeded up the latter toward 
Cerro Quia on the Colombian border. At the base of Cerro Quia, he 
found only one pair, below the outlet of the Rio Mono. 

A series of specimens throughout the range from Mexico to Panama 
shows no geographic variation. A form T. a. rhizophorae described by 
van Rossem (Proc. Biol. Soc. Washington, vol. 32, 1939, p. 155) on 
the basis of supposed greater amount of white in the frontal and loral 
area, from Tobari Bay, southern Sonora, ranging to Nayarit, proves 
invalid from the series now available. 

Hellmayr (Cat. Birds Amer., vol. 9, 1935, p. 70) has listed another 
swallow, Tachycineta stolgmanm Philippi, known only from Chepén 
on the coast of northern Peru, also as a race of albilinea, but this re- 
quires verification. In brief, this Peruvian bird originally was described 
as Hirundo leucopygia by Taczanowski (Proc. Zool. Soc. London, 
1880, p. 192) from 4 specimens collected by Stolzmann. As this name 
was preoccupied by an earlier H. leucopygia of 1834, Philippi (Ann. 
Mus. Nac. Chile, Zool., vol. 15, 1902, p. 23) renamed it Hirundo stolz- 
manni. The type, in the Warsaw Museum, is lost. Hellmayr, who 
examined one of the original specimens in the Senckenberg Museum in 
Frankfurt, noted that, compared to Hirundo albilinea of Central Amer- 
ica, it differed in much smaller bill, lack of white line above the lores, 
grayish instead of white bases on the dorsal feathers, gray instead of 
white underwing coverts, and grayish instead of white undersurface, 
with other minor differences. In spite of the extensive gap in the range, 
he listed the Peruvian population as a race of albilinea. In this he was 
followed by Peters (Check-list Birds World, vol. 9, 1960, p. 82). Hell- 
mayr’s action was accepted also by Zimmer (Amer. Mus. Nov., no. 
1723, 1955, p. 33). Since, in spite of modern studies in the area con- 
cerned, there is no other report of T. albilinea in northwestern South 
America, the relationship suggested appears doubtful. 7. albilinea, 
therefore, is listed as without accepted races. 


TACHYCINETA THALASSINA LEPIDA Mearns: Violet-green 
Swallow, Golondrina Verde 


Tachycineta lepida Mearns, Proc. Biol. Soc. Washington, vol. 15, March 5, 1902, 
p. 31. (Campbell’s Ranch, Laguna Mountains, Coast Range, 20 miles north of 
Campo, San Diego County, California.) 


12 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Medium size; undersurface white; above, green, glossed with purple, 
with white patches on either side of rump. 

Description—Length 120-130 mm. Adult male, crown and hind- 
neck bronze to purplish green, with a narrow greenish band across 
lower hindneck; back and scapulars definitely with a purplish tint but 
greener than crown; rump and upper tail coverts purple mixed with 
green; wings and tail dull black, with a bluish green sheen; undersur- 
face, including cheeks, scapulars, and undertail coverts, pure white, 
this color extending posteriorly over sides of rump; underwing coverts 
pale gray. 

Female, duller, greener, less purplish above; crown dull grayish 
brown, in some with a greenish sheen; side of head and upper hindneck 
grayish white; undersurface light grayish white on foreneck, upper 
breast, and scapulars. . 

Immature, dark grayish brown above; wings and tail dull black; 
sides of head and undersurface as in female. 

Measurements —Males (10 from Colorado, New Mexico, Arizona, 
California; April-July), wing 114.2-117.0 (115.9), tail 43.8-49.2 
(46.6), culmen from base 7.1-8.9 (7.8), tarsus 11.0-12.7 (11.5) mm. 

Females (10 from Colorado, New Mexico, Arizona, California; 
April-July, December), wing 108.2-114.8 (111.1), tail 41.7-46.3 
(43.3), culmen from base 7.1-8.6 (7.6), tarsus 10.5-13.9 (12.0) mm. 

Migrant from the north, rare (irregular). Known from numerous 
sight records in late February 1960, on the lower slopes of Volcan de 
Chiriqui, Boquete (El Salto), Volcan (El Hato), and Cerro Punta, 
western Chiriqui. 

Dr. Kugene Eisenmann, with Mr. and Mrs. James Linford of Oak- 
land, California, on February 21-22, 1960, found about 50 flying with 
Chestnut-collared Swifts near El Salto, above Boquete, Chirigui. From 
February 23 to 27, in late afternoon, from 70 to 100 were recorded 
daily over open pastures near the village of Cerro Punta. In the after- 
noon of February 27, near El Hato del Volcan, 1 perched on a wire, 
where Eisenmann observed it close at hand. 

The Violet-green Swallow nests from the Yukon Valley and Yukon, 
Canada, to northern Mexico. In winter it is found regularly to El 
Salvador and Honduras. Southward, in Costa Rica, specimens have 
been taken at Punta Piedra and Bebedero, Guanacaste, and Matina on 
the coast of Limon. 

In the winter of 1976-77 flocks were again seen in Panama. A. 
Greensmith reported seeing small numbers daily at Cerro Campana, 
western Panama Province, from December 27-29, 1976, with up to 40 


FAMILY HIRUNDINIDAE 13 


the last day, usually feeding with larger numbers of Blue-and-white 
Swallows. Then, up to 80 were seen feeding over open hillsides near 
the Volcan Lakes, western Chiriqui, on January 13-14, 1977. All field 
marks were noted with care, including the white patches on the sides of 
the rump, the white vent, and the white on the sides of the head (Tou- 
can, vol. 4, no. 2, 1977, pp. 6-7). 


HIRUNDO RUSTICA ERYTHROGASTER Boddaert: Barn Swallow, 
Golondrina Tijereta 


_Hirundo erythrogaster Boddaert, Table Planch. Enlum., 1783, p. 45. (French 
Guiana. ) 


Size medium; adult, in full plumage, with long, deeply forked tail 
(the lateral feathers often lost during the winter molt, so that this char- 
acter is not evident; identified then by the steel blue back and rump). 

Descripion.—Length 149-186 mm. Adult male, forehead chestnut; 
rest of upper surface steel blue, including edging on tertials and mid- 
dle wing coverts; wings and tail dusky; inner webs of rectrices (ex- 
cept central pair) with a white spot; malar area, chin, throat, and fore- 
neck to upper margin of chest cinnamon-rufous; steel blue color or 
lower side of neck extending over the side of the chest, in some the 
two sides joined; rest of undersurface, including sides, axillars, under- 
wing coverts and undertail coverts, pale cinnamon-rufous. 

Adult female, usually similar, but in some the undersurface paler. 

Immature, much duller colored; forehead dull white to buff; crown 
and hindneck dull brown to sooty black; undersurface duller, browner; 
in some, breast, sides, and abdomen white. 

Measurements —Male (10 from eastern and western United States, 
British Columbia, and Alaska), wing 114.3-123.6 (119.9), tail 71.2- 
92.8 (82.9), culmen from base 9.4-11.5 (10.7), tarsus 11.1-12.0 
(11.6) mm. ; 

Females (10 from eastern and western United States, British Co- 
lumbia, and Alaska), wing 109.1-119.2 (115.7), tail 69.7-83.3 (76.3), 
culmen from base 10.0-11.9 (10.9), tarsus 11.2-12.2 (11.6) mm. 

Migrant from the north, abundant; common winter resident espe- 
cially on the more open Pacific slope from early September through 
April, with many in passage to and from wintering grounds in South 
America. A few remain through the period of northern summer. Most 
common in the lowlands but ranging also across the higher levels. 

Migrant flocks en route to and from South America move regularly 
over the sea along the Caribbean coast, and also are seen crossing the 


14 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Gulf of Panama and other waters of the Pacific in the direction of 
Darién or Colombia. Thus, single birds or small groups of Barn Swal- 
lows may appear on any of the offshore islands. In stormy weather 
during periods of migration, hundreds may be encountered. Northward 
migration is evident by late March and continues into late May, when 
it becomes difficult to separate northern travelers from summering 
birds. Birds such as the 2 seen by Eisenmann at Coco Solo, Canal Zone, 
on July 23, 1956, could be summering or early transients moving south- 
ward. 

Migrant and wintering groups gather at night to sleep in low trees or 
reeds around lowland lagoons and marshes, spreading widely in their 
feeding activities during the day. The roosts are in the open. In 1966, 
at Puerto Armuelles, Chiriqui, the Chiriqui Land Company had built 
a roof over a long pier to allow handling of extensive shipments of 
bananas regardless of weather. During February and March, Barn 
Swallows gathered nightly to sleep under this shelter, undisturbed by 
the electric lights that illuminated the area throughout the night, or the 
workmen busy beneath them. 

The annual molt of adults and young comes during the months of 
northern winter. The full-plumaged birds seen regularly in early May 
are assumed to be those that will nest in the more northern parts of the 
breeding range in Alaska and Canada. 

Ridgely (im litt.) notes that in recent years Barn Swallows have in- 
creasingly come to be associated with sugarcane fields, especially dur- 
ing northern winter months; they hawk for insects over and around 
them, and also roost in them. This seems to be the pattern as far south 
as eastern Brazil. The considerable increase in Barn Swallows in the 
past decade may be associated with the great increase in sugar growing 
in Pacific slope lowlands of Panama. 


PETROCHELIDON PYRRHONOTA (Vieillot): Cliff Swallow, 
Golondrina de Paso 


Medium size. Tail short, slightly notched at tip; forehead white, buff 
or chestnut-brown; rest of crown black or dusky; rump buff to brown. 

Description Length 120-151 mm. Adult (sexes alike), crown and 
back shining blue-black; forehead white to chestnut (according to 
race); rump cinnamon-rufous; upper tail coverts grayish brown, edged 
with paler gray; wings and tail dusky brown; secondaries edged nar- 
rowly with white toward the tips; lores blackish; side of head and fore- 
neck chestnut, in some this color extending narrowly across hindneck; 


FAMILY HIRUNDINIDAE 15 


an irregular black mark variable in size, on foreneck, most prominent 
in adult males, in females often much restricted; sides more or less 
brownish gray; breast and abdomen white. 

Immature, colors much duller; feathers of the upper surface usually 
with paler margins, and color patterns as a whole less definite. 

Cliff Swallows are seen mainly in fall, and in the months of spring, 
from late February through April, usually in company with other mi- 
grant species of the family. It is probable that their principal winter 
quarters are in South America, so that most may be in transit when 
seen in Panama, but winter records have gradually accrued, suggesting 

that the species regularly winters in Panama, at least in small numbers. 

Geographic races in this species, widespread in its distribution in 
the breeding season, are not always clearly evident. On their northern 
breeding grounds 2 main groups may be recognized, one—in which the 
forehead is white—that is mainly northern, and the other—with this 
area chestnut-brown—found in the southern section. The distinction 
of subspecies in those of typical markings 1s clearly evident, but in the 
broad area where the two approach during the nesting season, the 
colors mentioned merge through intergrades so completely that there 
is no clear-cut division. As a whole, northern populations are larger, 
but here again, there is intergradation. From the considerable series of 
specimens available from Panama, 4 groups may be separated on the 
basis of color and size. Their records of occurrence are listed under the 
four headings that follow. (Two others that have been described are 
not clearly separable. ) 

While Cliff Swallows as a species may be identified readily in life, 
the various forms are so similar that the race may not be determined 
unless the bird is in the hand. Usually they are seen flying in company 
with other swallows, most frequently with the more abundant Barn 
Swallows. From available dates, the fall flight southward is recorded 
in Panama from August 21 through September to October 6. J. Karr 
and Ridgely (1m litt.) saw 2 at Chiva Chiva, Canal Zone, on July 29, 
1968. 

Winter records known to Ridgely (im litt.) include 1 at Tocumen, 
with 50 Barn Swallows on January 29, 1973; 6 at Flamenco Island 
(Fort Amador) on January 30, 1973; 2 at La Jagua on January 27, 
1976; and from Chiriqui, 1 at Las Lajas on February 13, 1976. 

Spring migration evidently begins early. Ridgely and R. McArthur 
saw 1 Cliff Swallow flying north over Panama Bay on February 12, 
1970. In March and April this species can be as abundant as Barn 
Swallows. On March 6, 1976, for example, Ridgely saw thousands 


16 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


along the Pan American Highway from David eastward for 30 km. 
Cliff Swallows are found in Panama as late as May 17, when in 1973 
Ridgely saw 50 near Escobal. 


PETROCHELIDON PYRRHONOTA PYRRHONOTA (Vieillot) 


Hirundo pyrrhonota Vieillot, Nouv. Dict. Hist. Nat, vol. 14, September 1817, 
Dp: D19: (Paraguay) 


Characters.—Forehead clear white in most individuals, size smaller; 
wing 107-112.8 mm. 

Measurements—Males (10 from eastern and western United States 
and southern Canada), wing 108.4-112.8 (109.8), tail 46.6-50.9 (48.9), 
culmen from base 8.9-10.6 (9.7), tarsus 11.3-13.2 (12.5) mm. 

Females (10 from eastern and western United States), wing 107.1- 
109.9 (108.4), tail 44.7-49.0 (47.9), culmen from base 9.4-10.2 (9.7), 
tarsus 12.1-12.8 (12.5) mm. 

Migrant from the north, common. 

Specimens examined are as follows: Male, in the British Museum 
(Natural History), received in the Tweedale Collection, without date 
of collection, purchased from the dealer G. A. Frank of London. It is 
assumed to be from the Canal Zone, as it shows the characteristic prepa- 
ration of McLeannan. One of his small labels attached reads “45 male.” 
The wing measures 110.5 mm. 

Immature male, USNM no. 30556, collected at sea ““N. of Panama, 
Oct. 20, 1863, Capt. J. M. Dow.” The locality is uncertain, but prob- 
ably means north of Panama City, off the west coast. The wing mea- 
sures 108.0 mm. 

Female, no. 2378, collection of Gorgas Memorial Laboratory, from 
Almirante, Bocas del Toro, October 10, 1960; wing 111.0 mm. 

Male aduit, USNM no. 461062, Las Lajas, Chiriqui, February 26, 
1956; wing 109.5 mm, A. Wetmore. 

Adult male, AMNH no. 500904, Panama City, April 3, 1897; wing 
MOS CN s Glarus: 

As the breeding range of this race extends widely through the central 
and northern United States, this subspecies may be the one most com- 
monly represented in the migrant flocks that come to Panama. These 
are seen commonly through September and October, and again from 
late I*ebruary through March to early May. Apparently the majority 
are birds of passage with the main wintering ground in South America. 

There is considerable variation in size in this nominate race, as indi- 
cated by wing length. Those that nest in the west, from Oregon north- 


FAMILY HIRUNDINIDAE 17 


ward, merge gradually into the large population (hypopolia) of Alaska. 
Intermediates from Oregon have been described as a distinct race, 
aprophata, with the “lower parts paler, and forehead more buffy (less 
clearly whitish)” (Petrochelidon albifrons aprophata Oberholser, Sci- 
entific Publications Cleveland Museum, vol. IV, no. 1, 1932, p. 6). In- 
dividual variation is such that this proposed additional race may not be 
distinguished with certainty, especially in winter migrants. 


PETROCHELIDON PYRRHONOTA HYPOPOLIA Oberholser 


Petrochelidon albifrons hypopolia Oberholser, Canadian Field-Nat., 1920, vol. 33, 
~ no. 5, p. 95. (Fort Norman, Northwest Territories.) 


Characters.—Forehead white; size larger. 

Measurements —Males (10 from Alaska and MacKenzie), wing 
112.7-116.1 (114.1), tail 45.7-51.7 (49.2), culmen from base 9.0-10.9 
(9.8), tarsus 12.4-14.2 (13.3) mm. 

Females (10 from Alaska and MacKenzie), wing 110.5-115.5 
(113.1), tail 47.1-53.9 (50.4), culmen from base 9.1-10.6 (9.7), tarsus 
12.4-15.0 (13.6) mm. 

Migrant from the north, abundance uncertain. At the Ciénaga Santo 
Domingo, eastern Province of Panama, below La Jagua, on March 30, 
1949, when a flock of 100 or more Cliff Swallows circled near, I shot 1 
and the others disappeared. The bird taken was extremely fat and also 
of maximum size, with the wing measuring 116.5 mm. It agrees fully 
in this with the population of northern Alaska and northwestern Can- 
ada, and is identified as the race nesting in that area. It is the only 
definite record of this subspecies from Panama that I have seen. 


PETROCHELIDON PYRRHONOTA TACHINA Oberholser 


Petrochelidon lunifrons tachina Oberholser, Proc. Biol. Soc. Washington, vol. 16, 
February 21, 1903, p. 15. (Langtry, Texas.) 


Characters.—Forehead light to fairly dark buff (varying from dull 
buffy white to dull brown). Size small. 

Measurements—Males (10 from Chihuahua, Guanajuato, and 
Texas), wing 100.0-105.1 (102.5), tail 42.8-45.7 (44.3) culmen from 
base 8.4-9.9 (9.3), tarsus 11.8-12.6 (12.0) mm. 

Females (10 from Texas and New Mexico), wing 101.3-104.7 
(103.0), tail 42.6-47.6 (45.2), culmen from base 8.4-9.9 (9.1), tarsus 
11.2-12.2 (11.9) mm. 

Migrant from the north. Abundance uncertain. 

On February 26, 1956, I collected a female, at Las Lajas, near the 


18 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


coast in eastern Chiriqui. The bird was with a mixed flock of the 
species, the others taken included birds assigned to melanogaster and 
pyrrhonota. 


PETROCHELIDON PYRRHONOTA MELANOGASTER (Swainson) 


Hirundo melanogaster Swainson, Philos. Mag., n.s. vol. 1, 1827, p. 366. (Real del 
Monte, Hidalgo, Mexico.) 


Characters—Forehead dark chestnut; size small. 

Measurements.—Males (10 from San Luis Potosi, Durango, Tepic, 
and Arizona), wing 101.4-106.2 (103.0), tail 42.4-46.5 (44.9), culmen 
from base 7.7-9.5) (8.5), tarsus, 105-127 ((113))) am 

Females (10 from San Luis Potosi, Durango, Sonora, Oaxaca, and 
Arizona), wing 101.4-109.2 (104.4), tail 40.6-45.6 (43.3), culmen from 
base 8.0-9.4 (8.7), tarsus 11.0-12.1 (11.5) mm. 

Migrant from the north. Apparently common. 

Specimens from Panama are as follows: A male and 2 females, col- 
lected February 26, 1956, on the Savannah Santa Cruz, near the sea, 
below Las Lajas, Chiriqui, where we encountered a small flight of Cliff 
Swallows. On March 20, 1961, at La Jagua, 3 small flocks circled over 
the savannas. When one group came near I collected a female of this 
race. There is also a male in the Museum of Comparative Zoology, 
taken by Wedel on October 6, 1930, at Puerto Obaldia, San Blas, (Gris- 
com, Bull. Mus. Comp. Zool., vol. 72, 1932, p. 358). 

Birds from northwestern Mexico and southwestern United States 
have been described as another race (Petrochelidon albifrons minima 
van Rossem and Hachisuka, Trans. San Diego Soc. Nat. Hist., vol. 9, 
no. 2, 1938, p. 5), which, however, from material seen is not separable 
from melanogaster. 


[PETROCHELIDON FULVA (Vieillot), Cave Swallow, 
Golondrina Pueblera 

A specimen of this species, no. 85.3.24.59, in the British Museum 
(Natural History), is labeled “Panama. McLeannan,” with no other 
data. It is in fair condition, except for the feathers of the throat and 
side of the head, which are considerably stained, perhaps by accident 
during preparation. It resembles the nominate population of fulva of 
the Greater Antilles. 

Salvin and Godman (Biol. Centr. Amer., Aves, vol. 1, 1883, p. 228) 
listed it as follows: “A single specimen, in bad condition, sent to us 
by M’Leannan from Panama, must ... be referred to P. fulva...As 


FAMILY HIRUNDINIDAE IQ 


M’Leannan once had in his possession a few Cuban birds which he 
obtained in exchange, it is just possible that his specimen of P. fulva 
may have come from Cuba, and not from Panama; but, on the other 
hand, one of the island birds may have strayed thus far.” The bird, in 
immature plumage, has no additional data in the museum catalog. The 
species differs from the Cliff Swallows in paler rufous of the head 
markings, darker chestnut of the rump, and lack of black spot on the 
throat. 

Sharpe (Cat. Birds Brit. Mus., vol. 10, 1885, addenda, p. 636) lists 
it under P. fulva as “n. Imm. sk. Panama (McLeannan). Salvin- 
Godman Coll.” Ridgway (Birds North and Middle Amer., pt. 3, 1904, 
p. 55) cited it as ““Panama?,” with references to Salvin and Godman, 
and Sharpe. 

One Cave Swallow was carefuily observed with other migrant swal- 
lows at Juan Diaz, eastern Province of Panama, on March 10, 1976, by 
Ridgely (in litt.). One was seen at Tocumen, eastern Province of Pan- 
ama, by V. Emanuel on February 9, 1980. 

The data are too uncertain to warrant inclusion of this species as a 
valid record. The winter range of the Cave Swallow is still unknown.| 


PROGNE CHALYBEA CHALYBEA (Gmelin): Gray-breasted 
Martin; Golondrina Urbana 


Hirundo chalybea Gmelin, Syst. Nat., vol. 1, pt. 2, 1789, p. 1026. (French Guiana.) 


Slightly smaller than the migrant Purple Martin; end of outer tail 
feathers somewhat narrowed and tip of tail notched; male and female 
alike in color pattern on undersurface. 

Description—Length 160-180 mm. Adult male, upper surface, in- 
cluding the lesser wing coverts, dark steel blue, changing on forehead 
to dull brown; wings and tail black, faintly bluish on outer webs; fore- 
head, side of head and neck, breast and sides dull brown, paler on upper 
throat which often is mottled with white; upper breast and sides edged 
more or less with white rarely with the white extensive; lower breast, 
abdomen, and undertail coverts white, with slightly marked narrow 
shaft lines of black on undertail coverts; axillars and underwing coverts 
dark sooty gray; longer underwing coverts usually tipped with white. 

Adult female, similar, but duller blue, more brownish above. 

Immature, distinctly browner above; upper foreneck and throat 
whitish in varying amount; breast duller, usually with paler tips on the 
feathers. 

A female, with ovaries not developed, taken at Olivo, 9 km northeast 


20 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


of Puerto Armuelles, Chiriqui, February 11, 1966, had the iris dark 
brown, bill black, tarsus and toes fuscous-black, and claws black. An- 
other female, a fully grown bird of the year, collected on the Rio 
Potrero, near the town of El Potrero, Coclé, March 7, 1962, had the 
iris dark brown, bill black, tarsus and toes fuscous-brown, and claws 
black. é 

Measurements ——Males (10 from Chiriqui, Veraguas, Herrera, 
Coclé, and Province of Panama), wing 126.9-134.9 (130.1), tail 60.8- 
65.8 (62.8), culmen from base 14.0-15.3 (14.7), tarsus 14.3-15.4 
(14.6) mm. 

Females (10 from Chiriqui, Veraguas, Coclé, Province of Panama, 
and Darién), wing 124.2-132.2 (128.0), tail 58.8-62.7 (60.8), culmen 
from base 14.5-15.5 (15.0), tarsus 14.0-15.0 (14.3) mm. 

Resident. Common and widely distributed throughout the main- 
land of the Republic, from the lowlands to 1500 m. In the Chiriqui 
highlands it has been recorded at Boquete and the Volcan area. Present 
regularly on Islas Coiba and Cébaco; less abundant, or casual after the 
breeding season, on islands in the Gulf of Panama: Isla San José 
(March 6-8, 1947), Brava. (In March 1904, W. W. Brown recorded 
a colony nesting on the church at San Miguel, Isla del Rey.) 

Possibly there is some seasonal movement among them, as in April, 
following heavy rains, I have seen hundreds resting on wires in the 
Savanna areas. 

The species is one of wide range from Mexico to southern South 
America. In Panama it is present regularly in the towns, especially 
around churches and other buildings in the plazas. In country districts 
it frequents open pasturelands and fields, where dead trees offer con- 
venient perches. It is usual to find a few associated in small groups 
from February to June during the nesting season. Following this, they 
gather in flocks that in evening come to communal roosts. Where these 
are located in trees in plazas, they may be a nuisance, and more so when 
they enter openings in the eaves and roofs of buildings. 

Their warbling calls are heard mainly around their nest sites, espe- 
cially in the period when they are pairing and building. On the whole, 
they are far less vociferous in the breeding season than the Purple 
Martin of North America at the same period. Nests are placed in cavi- 
ties in trees, frequently in holes made by woodpeckers. This original 
custom is varied when they settle around buildings, commonly nesting 
under eaves and hollow roof tiles or in holes in walls, building on beams 
or in recesses where space may be available. Nests are constructed of 
twigs, dried grasses, and other vegetable material, with string or bits of 


FAMILY HIRUNDINIDAE 21 


paper added when the site is in or near human habitation. In any lo- 
cation they seem less gregarious than the Purple Martin, as the nest 
sites occupied normally are separated, not close together. On Gatun 
Lake, Canal Zone, pairs build regularly in the top of the metal chan- 
nel markers. In the nesting period these martins are vigorous in attack- 
ing hawks and other large predatory birds that attempt to perch on or 
near their nesting trees. 

Eisenmann writes that the full extent of the nesting season in Pan- 
ama still remains to be determined: pairs with eggs or nestlings have 
been reported from March through early August, and he has seen pairs 
seemingly on territory near a tree hole or building cranny in February. 
On the San Blas coast and the nearby island of Mulatupo on Decem- 
ber 1 and 2, 1962, he saw a few isolated pairs about nest sites, sug- 
gesting that the nesting season may extend over much of the year, un- 
less pairs guard nest sites out of the nesting period. 

fiiieseses are white, with from 2 to 5 recorded in the set. Size 
ranges from 21.4-26.3 by 15.0-17.1 mm. (Meise in Schonwetter, 
Penmdoe@ol- part 17,1970, p. 193). A. Hartley (in Beebe, Hartley, 
and Howes, Tropical Wildlife in British Guiana, vol. 1, 1917, pp. 328- 
334), in an account of this species, recorded careful nest sanitation in 
which the parents removed the excrement of the nestlings. Food, as 
usual in this family, 1s composed of insects taken on the wing. Like 
other swallows, they swoop regularly to the surface of open waters to 
drink. Two collected by Burton (Bull. Brit. Orn. Club, 1975, p. 92) at 
Pacora, Province of Panama, weighed 40 and 44 g. 

At El Real, Darién, on January 7, 1964, I saw a curious aggressive 
reaction on the part of a Gray-breasted Martin that perched regularly 
for hours each day in a high dead treetop opposite our house. One after- 
noon, a skipper butterfly (Family Hesperiidae), in its usual rapid, er- 
ratic flight, darted about over the trees and houses. Its restless activity 
seemed to irritate the martin, as whenever the insect rose into the open 
air, the bird swooped at it and then returned to its perch. This con- 
tinued until finally the martin hit the insect with its breast, a clearly 
audible blow that knocked the skipper to the ground. There it lay for 
a minute or two with spread wings, partly stunned, until presently it 
rose, darted away, and disappeared. There was no indication that the 
bird had tried to seize it. 

Eisenmann believes that some birds of this race from Mexico at 
least occasionally winter or pass through Panama, as birds of northern 
Mexico are known to be migratory and flocks of this species are some- 
times seen when local birds are nesting. The southern race (domestica) 


22 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


of southern South America is highly migratory and regularly reaches 
Suriname, northern Brazil, and Venezuela; at least occasional occur- 
rence in Panama is not improbable during the austral winter but speci- 
men examination would be required. As in P. elegans, the molting 
primaries in July would be a strong clue that domestica was involved 
(see Eisenmann, Auk, 1959, pp. 528-532). 


PROGNE SUBIS (Linnaeus): Purple Martin, Golondrina Azul 


Largest of the swallows found in North and Central America (aver- 
aging larger than the Brown-chested Martin, Progne tapera). 

Description.—Length 170-200 mm. Adult male glossy dark steel 
blue above and below; partly concealed feather tufts on sides of lower 
back white; primaries dull black, with the basal area of the shafts dull 
white basally, changing to dull brown for the remainder of the length. 

Adult female, duller colored above (through partial exposure of the 
basal area of the feathers); forehead and more or less of forecrown 
sooty gray to white (varying racially and individually in depth of 
color); lores and auricular region dusky; sides of neck grayish, many 
individuals with an indefinite dull grayish band across the hindneck; 
anterior lower surface sooty gray, tipped narrowly with dull grayish 
white; lower breast, abdomen, and undertail coverts white, in some 
streaked indefinitely with sooty gray; sides, underwing coverts, and 
axillars dark sooty gray. 

Immature (both sexes), like female but duller above, with glossy 
area mainly on crown; wing coverts and secondaries tipped narrowly 
with dull grayish brown; collar across hindneck usually distinct. 

The Purple Martin is widely distributed in the nesting season from 
Baja California, and the mountain areas in the central plateau of 
Mexico; northward irregularly through the United States to Vancouver 
Island and southern British Columbia, and central eastern Canada. 
Following the breeding season, adult and immature martins gather at 
night in tree roosts that may include hundreds (even thousands) of 
individuals. At the end of August and in early September these groups 
disappear as the birds move southward to wintering grounds in South 
America. In spring when they return north they appear individually or 
in small groups on their nesting grounds. 

Two geographic races have long been recognized, the nominate form, 
Progne subis subis, being the breeding bird found widely through the 
United States north to south-central Canada. The slightly smaller and 
paler subspecies, Progne subis hesperia, is the nesting form of Baja 


FAMILY HIRUNDINIDAE 23 


California and a western montane race has been described by Behle 
(Condor, 1968, p. 166) as Progne subis arboricola, with the type lo- 
cality at ‘Payson Lakes, 8300 feet elevation, 12 miles southeast Payson, 
Utah County, Utah.” The assigned nesting range is from the Wasatch 
and Pavant Mountains of Utah, south to Arizona from the Mogollon 
Plateau to the Chiricahua Mountains. It is described as larger than 
subis, with the female like hesperia in paler color. Wing measurements 
of specimens that I have collected in northwestern Utah are as follows: 
Males, 152.0-157.0 (154.7); females, 144.0-152.0 (148.0) mm. 

In fall migration, Paynter (Auk, 1953, p. 347), at the end of August 
and the beginning of September 1952, recorded Purple Martins in 
small number in southward migration through the Caribbean islets on 
the Campeche Bank north of the Yucatan Peninsula. Russell (A. O. 
U. Orn. Mon., no. 1, 1964, p. 130) noted them as “regular transients” 
through British Honduras in spring and fall. Monroe (A. O. U. Orn. 
Mon. no. 7, 1968, p. 278) found them fairly common in spring and fall 
on the Caribbean side of Honduras. From October 8 to 12, he recorded 
several males in the Choluteca-Namasigte region of the Pacific Coast. 
Martins are passage migrants spring and fall through Panama, but as 
yet in detail are relatively little known. 

The first two subspecies are recorded in Panama from specimens. 
The third may be found also among the migrants, since Purple Martins 
as a species pass the period of northern winter in Brazil. 

Dr. Eugene Eisenmann has furnished the following summary of 
occurrence in Panama from his personal observations and those of 
other observers sent him. The records are based partly on female and 
immature individuals, which in life differ from the resident Gray- 
breasted Martin (Progne chalybea chalybea) in larger size, and in 
pale area on the side of the neck and often on the forehead. In the 
Caribbean area of the Isthmus, in southward flight this martin has 
been recorded both in small groups and in flocks of 40 to 5000 birds, 
from August 3 to September 15, at Almirante, Bocas del Toro (N. G. 
Smith), near Coco Solo, Coco Solito, and Gatun Lake near Gamboa, 
August 5 and September 21 (FE. S. Morton), Barro Colorado Island 
(Willis), Gatun Dam, Canal Zone; Puerto Pilon, Colon (Ambrose). 
Reports from the Pacific slope are few: Anton and Penomomé, Coclé, 
September 18 (H. Loftin, E. Tyson), between Tocuman and Plata- 
nales, eastern Province of Panama, September 10 ( Pujals et al.) , south- 
western Canal Zone, one September 12 (R. Ryan). 

The spring migration northward is little known. At Almirante, 
Bocas del Toro, on February 18, 1958, during a steady rain, between 


24 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


35 and 40 Purple Martins came to perch on dead limbs in the top 
of a tall tree beside our living quarters. Later, on March 2, during 
travel by launch along the seaward side of the Valiente Peninsula, I 
noted others flying northward low over the water. Until March 6, I 
recorded them daily, resting in dead trees over our house, in flight 
crossing Almirante Bay, or offshore at Boca del Drago. Between 
March 19-21, 1962, Charles O. Handley, Jr., at vaniousocalimecwen 
route from Colon to Isla Escudo de Veraguas, recorded several flying 
northwest offshore over the Caribbean (Wetmore, Smiths. Misc. Coll, 
no. 145, 1963, p.2). An “all dark” Progne seen at Gamboa on April 1, 
1968, by Ridgely was most probably this species, while 2 such at Bayano 
Lake on April 24, 1968, with Brown-chested Martins, could be late 
transients or Southern Martins on their migration. 


PROGNE SUBIS SUBIS (Linnaeus ) 
Hirundo subis Linnaeus, Syst. Nat., ed. 10, 1758, p. 192. (Northeastern Manitoba. ) 


Characters—Size intermediate between P. s. hesperia and P.. s. ar- 
boricola; female with forehead and forecrown dark gray. 

Measurements.—Males (10 from the breeding range), wing 145.2- 
150.3 (146.6), tail 71.2-78.4 (74.2) culmen from base 13.1-14.8 (13.8), 
tarsus 15.0-15.7 (15.3) mm. 

I*emales (10 from the breeding range), wing 141.0-146.5 (144.5), 
tail 66.2-73.6 (70.1), culmen from base 14.2-15.4 (14.9), tarsus 15.2- 
1.529) (Ol 523))) sacien. 

Migrant and winter visitor from the north. This is the most widely 
distributed of the races. It is known in Panama definitely from the fol- 
lowing specimens: Coco Solito, Canal Zone, male, collected August 5, 
1955; female, August 3, 1956 (J. E. Ambrose, specimens in American 
Museum of Natural History); Almirante, Bocas del Toro, 2 males, 1 
female, February 18, 1958 (A. Wetmore, in the Smithsonian collec- 
tions ). 

The type locality for the species given by Linnaeus (from Edwards) 
as “ad sinum Hudsonis,’ Hudson Bay, is so listed in the A. O. U. 
Checklist (1957, p. 365). L.L. Snyder’ (Can: Field-Nat ave 77 ae am 
pp. 128-129) reports that “Edwards’ pre-Linnaean descriptions and 
illustrations were based on specimens collected by James Isham be- 
tween the years 1732 and 1745. ... When Isham lived in the New 
World he was stationed mostly at York Factory, but was briefly at 
Prince of Wales Fort (=Churchill).”’ From this data Snyder cites the 
type locality as “northeastern Manitoba.” 


FAMILY HIRUNDINIDAE 25 


PROGNE SUBIS HESPERIA Brewster 


Progne subis hesperia Brewster, Auk, vol. 6, no. 2, April 1889, p. 92. (Sierra de la 
Laguna, Baja California, Mexico.) 


Characters —Adult male, like nominate subis but smaller; female 
and immature male averaging paler in general coloration; foreneck, 
breast, and sides more extensive, white to pale grayish white. 

Measurements.—Males (10 from Baja California) , wing 140.5-145.0 
(142.2), tail 65.0-70.3 (67.7), culmen from base 13.4-15.9 (14.8), 
tarsus 14.9-15.9 (15.2) mm. 

Females (6 from Baja California), wing 136.1-140.7 (138.2), tail 
62.1-68.0 (66.6), culmen from base 13.5-15.6 (14.8), tarsus 14.8-15.5 
(15.1) mm. 

Passage migrant to and from winter quarters in South America. 
Known definitely in Panama from a male in immature plumage in the 
American Museum of Natural History, collected by R. R. Benson, at 
Cocoplum, Bocas del Toro, October 27, 1927. 

Zimmer (Amer. Mus. Nov., no. 1723, 1955, pp. 4, 5) listed this 
specimen, but with information available at the time was uncertain as 
to the characters of hesperia. The bird, on examination, is an immature 
male in good plumage, with the paler coloration of hesperia, evidently 
a bird from the preceding summer. With a wing measuring 143.6 mm, 
it is within the size limits accepted for this race. 

An early report of this race in migration is that of Richmond (Proc. 
U.S. Nat. Mus., vol. 16, 1893, p. 485). He collected a pair of Purple 
Martins September 13, 1892, in eastern Nicaragua, on a banana plan- 
tation on the Rio Escondido 50 miles above Bluefields, from ‘“‘a flock 
of six or eight which had settled in the top of a dead tree during a 
shower.” These 2 specimens (in the Smithsonian) belong to the sub- 
species hesperia and mark the first record for migrants of this form. 
Wing measurement for the male is 144.1 mm, for the female, 133.6 mm. 


PROGNE ELEGANS Baird: Southern Martin, Golondrina Negra 


Progne elegans Baird, Rev. Amer. Birds, May 1865, pp. 274 (in Key), 275. (Ver- 
mejo River, Paraquay = Rio Bermejo, Argentina.) 


Closely similar in appearance to Progne subis, but on average 
slightly smaller; in adult male blue sheen faintly duller, less brilliant; 
female more heavily marked below and lacking pale forehead and sides 
of neck; tail more deeply forked than in subis. 

Description.—Length 175-185 mm. Male, with concealed white on 
sides and flanks like swbis, but with undertail coverts duller, less clearly 


26 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


white; breast and lower undersurface much more heavily marked with 
dusky, lacking pale area on sides of neck and forehead. Immature male 
like female but with more blue sheen above and often on sides. 

Measurements.—Males (10 from Brazil, Argentina, and Uruguay), 
wing 137.8-142.5 (139.9), tail 60.5-82.8 (73.7), culmen from base 
12.2-15.4 (14.0), tarsus 15.4-17.8 (16.5) mm. 

Iemales (10 from Brazil, Paraguay, and Argentina), wing 132.0- 
14.0 (135.2) tail 65.7-77.8 (71.2), culmen from base 12.0-15.4 (13.8), 
tarsus 15.0-16.7 (16.0) mm. 

Casual; one definite occurrence in eastern San Blas. Breeds from 
the highlands of Bolivia and Argentina south to Chubut; in the austral 
winter regularly north to northern Brazil and Suriname. 

The single record for Panama is a specimen (in the University of 
Michigan Museum of Zoology) from Obaldia (Puerto Obaldia), San 
Blas, collected July 14, 1931, by Wedel, with sex marked “2? ONE” 
(apparently in error). The bird is immature, with longer wing and tail 
feathers much worn, in partial molt to adult stage. A few scattered 
feathers on the left side of the crown above the eye, and others in the 
tertials and wing coverts, are those of the dark blue male plumage. 
Though on casual glance the specimen resembles the female sex, on 
close scrutiny it appears to be a male beginning the change from juve- 
nile dress to that of the adult. (For the record of this specimen, see 
Eisenmann, Auk, 1959, pp. 528-532.) It is identified as elegans, as 
would be expected. 

While Peters (Checkl. Birds World, vol. 9, 1960, p. 88, following 
Hellmayr, 1935) includes this bird of eastern South America as con- 
specific with Progne modesta of the Galapagos, the latter form in series 
is much smaller, and also lacks the partly concealed white of sides and 
flanks found in P. subis and P. elegans. Progne elegans here is given 
separate specific status. 

The form murphyi, named by Chapman (Amer. Mus. Nov., no. 
187, 1925, p. 6), from the coast of Peru, possibly may connect the two, 
but this needs further checking (the Smithsonian at present has no 
specimens ). 

There have been observations in June and July of 1958 and 1970 
(Ridgely, 1976, p. 262) and on April 24, 1976 (Ridgely, in litt.) of all 
dark Progne in central or eastern Panama, dates when the northern 
migratory P. subis is very unlikely and when the southern migratory 
P. elegans may well occur (see under P. subis). Although separation 
of adult males in the field seems impracticable, females and immatures 
are often distinguishable, and some reports of probable adult males of 


FAMILY HIRUNDINIDAE 27 


P. elegans include observations of accompanying Progne that had 
characters of females or immatures of P. elegans. Eisenmann believes 
P. elegans to be an irregular migrant in small numbers to Panama. 


PROGNE TAPERA FUSCA (Vieillot): Brown-chested Martin; 
Golondrina Parda 


Hirundo fusca Vieillot, Nouv. Dict. Hist. Nat., nouv. ed., vol. 14, September 1817, 

D210. (Paraguay.) 

One of the larger swallows; white on throat and abdomen; a dark 
band across the breast; a few blackish spots in center of breast and 
abdomen. 

Description.—Length 155-170 mm. Adult (sexes alike), above dull 
dark brown, including side of head and wing coverts, the feathers 
edged indistinctly with lighter brown; wings and tail somewhat blacker; 
secondaries tipped narrowly with white; throat, lower breast, abdo- 
men, and undertail coverts white; sides and a broad band across upper 
breast dark brown; a series of spots on the tips of the lower central 
breast feathers dark brown to nearly black. 

A male of this race collected at Kilometer 80, west of Puerto Pinasco, 
Paraguay, September 18, 1920, had the bill, tarsus, toes, and claws 
black; iris bone brown. 

Measurements.—Males (10 from Argentina, Paraguay, Brazil, and 
Colombia), wing 134.6-143.0 (139.2), tail 64.4-68.0 (65.9), culmen 
from base 13.9-15.2 (14.6), tarsus 13.4-15.1 (14.6) mm. 

Females (10 from Argentina and Uruguay), wing 131.0-139.9 
(134.9), tail 61.8-66.5 (64.0), culmen from base 13.3-15.6 (14.5), 
tarsus 14.0-14.9 (14.4) mm. 

The winter migrants that come to Panama often have the wings and 
tail worn, or are in renewal after molt, so that their measurements may 
differ from those listed above. 

Austral winter migrant from South America, where it nests from 
eastern Bolivia and central Brazil south to northern Argentina, Para- 
guay, and southern Uruguay. Recorded from April to October in the 
Canal Zone and the savanna region of eastern Province of Panama. 
This southern form of the Brown-chested Martin, long recorded as a 
winter visitor to northern South America, now regularly extends its 
migratory flights to Panama. The species was first reported for Pan- 
ama by Eugene Eisenmann (Smithsonian Misc. Coll., vol. 117, 1952, 
p. 46) from 3 recorded at the Barro Colorado Island laboratory, Canal 
Zone, July 4, 1949. In July, 1949 and 1951, he observed many about 
the city of Panama, and in the Canal Zone. Since then the species has 


28 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


been reported regularly, although numbers vary greatly from year to 
year. The earliest specimens known from the Republic are 2 males 
collected by R. R. Benson near Tocumen, eastern Province of Panama, 
August 21, 1931 (received by the Smithsonian in a collection presented 
in 1954). 

The main group of migrants appears to arrive in the last week of 
April or in early May (with one seen by me April 5, 1954, at Pedro 
Miguel, Canal Zone, as an early date). Sometimes in late April flocks 
of hundreds can be seen in the Caribbean side of the Canal Zone. The 
birds are present in numbers through September, with a few remaining 
later (18 seen October 26, 1955, by J. E. Ambrose). They have been 
recorded most commonly in open areas of the Canal Zone, and the 
adjacent Province of Panama and also west to western Chiriqui. At 
evening they may join Gray-breasted Martins to spend the night in 
communal roosts. 

Eisenmann reports seeing them commonly during June, July, and 
August in some years on the Pacific slope of western Panama, includ- 
ing Coclé (Penonomeé, Anton), Veraguas (near Santiago), Chiriqui 
(Remedios, David, and many other lowland localities). On the Carib- 
bean slope he observed several as far west as Changuinola, Bocas del 
Toro, on June 30, 1956 (N. G. Smith noted 3 or 4 there on September 
3, 1964) and as far east as Rio Piedras, Colon. There seem to be no 
reports from the Azuero Peninsula, San Blas, or Darién, but this prob- 
ably indicates absence of observers in open areas during the season of 
occurrence. 

In August 1954 Eisenmann found them exceptionally abundant in 
open areas—greatly exceeding in numbers the aggregate of all other 
swallows seen; flocks of several hundreds often perched together on 
wires, and thousands had been noted in one day (August 9 in Coclé). 
By early September, numbers were much reduced, although Major 
Chapelle noted as many as 80 on September 18 on the Pacific slope of 
the Canal Zone, but numbers kept dropping until he saw his last Brown- 
chested Martin that year on October 16 (a lone bird perched with 
Gray-breasted Martins). Although Eisenmann has observed this 
species in Panama (since he first recognized it in 1949) whenever he 
has visited that country from May through early September, he has 
never seen numbers approaching those in 1954. In 1972 (August 2-4) 
Ridgely reports seeing flocks of hundreds around FE! Llano near the 
Bayano River in eastern Province of Panama. He saw 5 on January 
2, 1974, west of Rio Hato, Coclé—evidently birds that had failed to 
migrate to the breeding grounds. 


FAMILY HIRUNDINIDAE 29 


The brown upper surface and the band across the breast in this mar- 
tin present much the same color pattern as that of the much smaller 
Bank Swallow. In their winter flocks these martins are active in cir- 
cling in the air over wire lines and other perches. In the main, the 
migrants are silent. On their nesting grounds in Argentina | have heard 
them giving low calls, chu chu chup, that have little carrying power. 

Baird (Rev. Amer. Birds, 1865, pp. 272, 383) separated this species 
under the generic name Phaeoprogne, distinguished from Progne by 
slightly forked tail, with the tips of the feathers rounded, and weaker 
bill and legs, with a line of feathers on the inner margin of the tarsus 
for two-thirds its length. Sharpe (Cat. Birds Brit. Mus., vol. 10, 1885, 
p. 172) cited “P. tapera” as the type of Phaeoprogne. Zimmer (Amer. 
Mus. Nov., no. 1723, 1955, pp. 9-10) recognized the genus as valid, 
with two forms listed as races under tapera, the older of two names. 
The characters on which the genus Phaeoprogne was proposed by 
Baird are slight, so that it was not recognized as distinct from Progne 
by Mayr in his revision of Peters’s account of the Hirundinidae 
(Check-list Birds of the World, vol. 9, 1960, p. 85). 

The nominate race, Progne tapera tapera (Linnaeus), has the throat 
less extensively white, with the gray of the breast extending on the 
lower foreneck, and without the dusky black spots down the center of 
the lower breast and upper abdomen. It appears to be resident from the 
Caribbean coast and the Magdalena Valley, Colombia, to the Guianas 
and south to Peru and northern Brazil. 


NEOCHELIDON TIBIALIS MINIMA Chapman: White-thighed Swallow, 
Golondrina Muslos Blancos 


FIGURE 2 


Neochelidon tibialis minimus Chapman, Amer. Mus. Nov., no. 138, October 18, 
1924, p. 9. (Juntas de Tamana, Choco, Colombia.) 


Small; upper surface wings and tail dull black; underneath dull 
brown; feathered area of lower leg white. 

Description.—Adult (sexes alike), upper surface of body dull, some- 
what brownish, black; loral area, wings, and tail dull black; undersur- 
face, including underwing coverts, grayish brown; feathers of lower 
leg (slightly developed as a tuft) white. 

Immature, feathers of lower surface faintly edged with dull white. 

A male, collected at the head of the Rio Guabal, Coclé, July 28, 1962, 
had the iris dark brown; bill black; tarsus and toes fuscous-black; 
claws black. A female near the Peluca Hydrographic Station, Coldn, 


30 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


February 23, 1961, had the iris dark mouse brown; maxilla and tip of 
mandible black; base of mandible fuscous-black; tarsus and toes 
fuscous-brown; claws black. 

Measurements—Males (10 from northern Coclé, eastern Province 
of Panama, San Blas, and Darién), wing 80.5-88.5 (84.8), tail 37.5- 
45.8 (40.5), culmen from base 7.1-8.4 (7.7), tarsus 9.2-11.0 (10.1) 
mm. 

Females (10 from eastern Colon, Darién, and San Blas), wing 80.5- 
88.1 (85.1), tail 35.7-46.4 (41.4), culmen from base 7.2-10.3 (8.2), 
tarsus 9.0-10.5 (9.9) mm. 


Ficure 2.—White-thighed Swallow, Golondrina Muslos Blancos, Neochelidon 
tibialis minima. 


Resident. Found locally in pairs or small flocks in humid forest 
areas of central and eastern Panama; on the Pacific slope, in the high 
levels of Cerro Azul, especially around the lake; and in Darien, on the 
lower Rio Sambu, the Rio Tuira at the mouth of the Paya, at Cana, 
along the Rio Pucro to the village of Pucro; and around Cerro Tacar- 
cuna from the base to 1250 m; on the Caribbean side, in northern Cocleé; 
in the Canal Zone near the lower Rio Chagres; and eastern San Blas 
near Armila. 

On February 28, 1962, I collected a male from a small flock near 
the head of the Rio Guabal, northern Coclé. Ridgely (1976, p. 262) 
saw 6 at El Valle, Coclé, April 27, 1969. On January 13, 1976, J. J. 


FAMILY HIRUNDINIDAE 31 


Pujals observed 1 above Santa Fé, Veraguas. In the Canal Zone it 
may be found daily along the Pipeline Road, in the Fort Sherman-San 
Lorenzo area, and along the Achiote Road. 

In 1961, in the region behind Madden Lake, these swallows were 
fairly common along the Rio Boqueron in February, and on the Rio 
Pequeni in March. On Cerro Azul, C. O. Handley, Jr., found them 
numerous in June 1957, and the following year R. S. Crossin collected 
1 at the lake on August 12. Near Armila, eastern San Blas, I found 
them rather common in late February and early March at the edge of 
forest, back of the Indian farms. 

In the Tuira Basin, Darién, in March 1959, I saw a flock of 30 near 
the mouth of the Rio Paya. In late January and February 1964, they 
were fairly prevalent near the village of Pucro on the Rio Pucro; and 
on March 2, we caught 3 in a mist net set at 250 m on the north fork of 
this river, in the higher levels of Cerro Tacarcuna. Others were seen 
in March, lower down at the old village site near the base of the moun- 
tain. 

Distinguished always by tiny size, they ranged in small flocks, cours- 
ing over clearings or above the broader, open stretches of streams. 
Their flight often seems flitting and irregular compared to that of 
larger swallows with which they associate. They sometimes hawk for 
insects over the canopy with Chaetura swifts. Occasionally, I noticed 
them bathing by dipping to the surface of open stretches of water as 
they circled on the wing. At rest they gather separately from larger 
swallows, on small branches of dead trees above water or at the border 
of small openings in the forest. In my experience they were silent. 
Specimens collected from January to April seemed to be in a resting 
stage after the nesting season, as some were molting slightly over the 
body. 

A female taken by E. A. Goldman at Cana had the stomach crammed 
with fragments of tiny beetles, hemiptera, and hymenoptera.. Two col- 
lected in the Canal Zone area (Strauch, Bull. Brit. Orn. Club, 1977, 
p. 64) weighed 9.1 and 9.2 g. 

The only note on nesting seen is that of Griscom (Amer. Mus. Nov. 
no. 282, 1927, p.7). At the end of February 1927, when he was travel- 
ing by yacht, “two or three pairs were found on the Sambu River nest- 
ing in holes in the river bank directly under Indian huts. The birds 
were remarkably tame and confiding.” No account of the nest and eggs 
is known to me. Eisenmann, however, has seen pairs guarding wood- 
pecker holes in slender dead trees from at least early April to June, and 
has never noted nesting in banks. On September 14, 1965, in a wooded 


32 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


ridge near Cerro Azul he saw a full grown (flying) juvenal being fed 
by an adult; by this period many birds were in small flocks. 


NOTIOCHELIDON CYANOLEUCA (Vieillot): Blue-and-white Swallow, 
Golondrina Barranquera Azul 

Small; undersurface from throat to abdomen white; above, dark 
steel blue. 

Description —Length 110-125 mm. Adult (sexes alike), tail mod- 
erately forked; dorsal surface, including the scapulars, iridescent dark 
steel blue; lores and space immediately behind eye black; wings and tail 
dull black; undersurface with throat, breast, and abdomen white; up- 
per sides black; undertail coverts black (nominate race cyanoleuca), or 
black and white (patagonica). 

Immature, upper surface dull sooty brown on crown and back; else- 
where, including wings and tail, dull brown; throat, band across breast, 
and sides wood brown; throat, breast, and abdomen white; undertail 
coverts dull brown, or mixed dull brown and white. 

The dark (or partly dark) undertail coverts distinguish this species 
in life from the Mangrove Swallow, which also is small, with clear 
white undersurface, but with this color including also all of the under- 
tail coverts. In addition, the Mangrove Swallow has a white rump and 
a short, narrow white supraloral, lacking in cyanoleuca. 

Two geographic races are found in Panama. The nominate form 
breeds in Panama, and in South America, from Colombia to southern 
Brazil, Uruguay, and northern Argentina. The more southern breed- 
ing form, N. c. patagonica, is recorded as nesting from the mountains 
in northern Argentina to northern Tierra del Fuego. In the period of 
southern winter, patagonica moves northward, and with it move the 
more southern populations of nominate cyanoleuca. It is probable that 
the flocks recorded irregularly in Panama during the breeding season 
comprise migrants of both races. 


NOTIOCHELIDON CYANOLEUCA CYANOLEUCA (Vieillot) 


Firundo cyanoleuca Vieillot, Nouv. Dict. Hist. Nat., nouv. ed., vol. 14, September 
1817, p. 509. (Paraguay. ) 


Characters.—Adult, with all of the undertail coverts black; under- 
wing coverts dark to blackish grayish brown. 

Immature, undertail coverts dark brown, in some individuals tipped 
lightly with white; underwing coverts dark brown. 

An adult female, collected at 1350 m along the Rio Chiriqui Viejo, 


FAMILY HIRUNDINIDAE 35 


beyond El Volcan, Chiriqui, March 21, 1954, had the iris dark brown; 
tarsus, toes, and claws fuscous. Another adult female, taken at 1900 m 
on the west base of Volcan de Chiriqui, March 4, 1965, also had the iris 
dark brown, tarsus and toes dark brown, claws fuscous-black. 

Measurements.—Males (10 from western Panama and Costa Rica), 
wing 92.0-99.0 (94.3), tail 45.4-52.6 (49.0), culmen from base 6.7-8.0 
(7.2), tarsus 9.1-10.6 (9.8) mm. 

Females (10 from western Panama and Costa Rica), wing 87.5-97.1 
(91.5), tail 41.8-52.6 (48.0), culmen from base 6.0-7.9 (6.8), tarsus 
9.2-10.6 (9.9) mm. 

Resident. Common over open areas in the highlands of Chiriqui 
above 1000 m elevation, ranging to the higher levels; reported on upper 
Rio Chiriqui (Fortuna Dam site), central Chiriqui (Ridgely) re- 
corded on Cerro Flores, eastern Chiriqui, and also in the district of 
Calovévora, northern Veraguas; casual at Gatun, Canal Zone. It is 
also common at Cerro Campana, western Province of Panama, where 
Ridgely first found it in December 1967; it was then already numerous 
and had probably been there for some time. 

There is a specimen in the American Museum, taken by Griscom’s 
party east of the main mountain range in western Chiriqui on Cerro 
Flores, north of Remedios. A male and 2 females in the British Mu- 
seum, secured by Arce, are labeled Calovévora, northern Veraguas, 
1869. 

These small swallows, common over open areas, come regularly 
about houses and farm buildings. In several seasons I found them nest- 
ing on ledges and under the eaves of the house that we occupied at Palo 
Santo beyond El Volcan, Chiriqui. Hundreds also nested on rock cliffs 
on the high mountain slopes to 2800 m on El Bart. Nests were made of 
dried grasses and other vegetation, filling the crevices that they occu- 
pied. Some of the structures seemed loosely built, as on several oc- 
casions | found broken eggs that had fallen beneath the nests. 

Skutch (Auk, 1952, pp. 393-405, and Pac. Coast Avif. no. 34, 1960, 
pp. 279-280), in detailed studies in southwestern Costa Rica, found 
them using any available cavities, from those in rotted dead tree trunks 
to artificial shelters under the eaves of houses, crevices in walls, or 
occasionally tunnels dug by animals in banks of earth along trails and 
highways. The thatched roofs on houses and other buildings in country 
areas also furnish shelter. He reported sets of 2, 3, and 4 eggs. These 
were glossy white, without markings. Measurements of 6 eggs were 
16.3-16.7 X 11.5-12.2 mm. 

He recorded that laying begins in March, with both male and female 


34 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


incubating and caring for the young. At night, one parent slept on the 
eggs while the other rested on the rim of the nest. The newly hatched 
young have a sparse covering of light gray down. A second brood may 
come in June. When mated, the pair remains together throughout the 
year following the breeding season. 

In most of their activities, I have noted that these swallows are 
mainly silent. The song is described as “a thin, weak, long-continued 
trill which slides upward at the end.” 

One egg in the British Museum collection that I examined, taken by 
Underwood on Volcan Irazu, Costa Rica, April 25, 1901, was white, 
without markings and measured 16.9X12.6 mm. The maximum sizes 
(listed by Meise in Schonwetter, Handb. Ool., pt. 17, 1970, p. 193) 
which range larger, to 20 x 14 mm, in part may include specimens from 
South American localities where nominate cyanoleuca is not found. 

Fleas parasitic on birds are little known, so it is of interest to report 
one captured on a specimen of this northern race of the Blue-and-white 
Swallow, in March 1962, by Vernon J. Tipton and C. L. Hayward. 
In a report on the fleas (Siphonaptera) of Panama by Tipton and 
Eustorgio Mendez (in Wenzel and Tipton, Ectoparasites of Panama, 
Field Museum of Natural History, 1966, p. 310) this parasite is listed 
as Dasypsyllus sasius veneguelensis (1. Fox and Anduze), with the 
comment that it was abundant in swallows’ nests, both in buildings and 
in earth banks. “In one hole in an earth bank the fleas were very 
numerous in the sand below the scanty nesting material.” 


NOTIOCHELIDON CYANOLEUCA PATAGONICA 
(D’Orbigny and Lafresnaye ) 


Hirundo patagonica D’Orbigny and Lafresnaye, Syn. Av., in Rev. Zool., ann. 7, 

1837, p. 69. ( Patagonia.) 

Characters.—Adult, with only the lower half of the undertail coverts 
black, the upper area being white like the rest of the undersurface; 
underwing coverts paler grayish brown. 

Immature, with underwing coverts much paler gray; undertail co- 
verts paler with tips more extensively white. 

Measurements.—Males (10 from Darién, Argentina, Peru, and Ven- 
ezuela), wing 101.2-111.9 (104.5), tail 50.1-55.6 (52.6), culmen from 
base 7.1-8.5 (7.8), tarsus 11.1-12.4 (11.8) mm. 

Females (10 from Darién, Argentina, and Chile), wing 99.2-105.6 
(102.8), tail 44.7-57.1 (48.7), culmen from base 7.5-9.5 (8.2), tarsus 
11.4-12.6 (12.0) mm. 

Migrant from southern South America; of irregular occurrence. 


FAMILY HIRUNDINIDAE 35 


The first published record for Panama is by Chapman (Bull. Amer. 
Mus. Nat. Hist., vol. 55, 1926, p. 558) of an adult male in the Prince- 
ton Museum collected by “J. P. Chapin at Juan Mina, Rio Chagres, 
Panama, July 17, 1923, while a member of the Charles Rogers Prince- 
ton Expedition.” In the British Museum there are 4 earlier specimens 
from Panama—3 immature birds marked as taken by McLeannan and 
the fourth, an adult, though not labeled, from its style of preparation 
evidently also from this collector. One of the immature individuals is 
marked Lion Hill Station, the others are indicated only as from Pan- 
ama. It is probable that these were sent by the collector to Salvin fol- 
lowing McLeannan’s visit to Panama in 1873, as they were received in 
the Salvin and Godman collection, the 3 immature birds having been 
presented in 1885, the adult in 1888. 

A male and female in the Academy of Natural Sciences collected by 
Jewel on July 7 and 16, 1911, from flocks at Gatun, Canal Zone, were 
attributed to the nominate form, but proved to be undoubtedly of the 
migratory southern race patagonica when reexamined by James Bond 
and R. Meyer de Schauensee (Bond to Fisenmann, im litt.). 

At Cana, on Cerro Pirre, Darién, on May 22, 1912, E. A. Goldman 
recorded “‘a flock containing hundreds seen perched on wires. Later in 
the day at least 100 were sitting close together on a spot where the 
ground was bare, and where I was unable to find anything to attract 
them.” The 2 specimens that he collected are of the present race. 

On April 5, 1954, I saw a swallow resting on high wires opposite the 
fire station at Pedro Miguel, Canal Zone, that appeared definitely to be 
patagonica. From its elevated position facing me, I had clear view of 
the undertail coverts, which showed the restriction of the black color 
to the lower area that marks this subspecies. Guy Tudor saw one over 
the Bayano River below Chepo on April 2, 1971 (Ridgely, im litt.). 

Farther north, migrant individuals of patagonica have been collected 
from groups of other swallows by Howell, July 9, 1954, at El Recreo, 
Depto. Zelaya, southeastern Nicaragua (Condor, 1955, p. 188), and by 
Alvarez del Toro, May 24, 1954, at Tuxtla Gutierrez, Chiapas, Mexico 
(Condor, 1957, p. 268). 

On June 17, 1971, Eisenmann, with J. J. Pujals, carefully examined 
a group of swallows on a lowland roadside telegraph wire between 
Chepo and La Capetana, eastern Province of Panama, that included 1 
South American migrant P. tapera fusca, T. albilinea albilinea, and 11 
N. c. patagonica. Ridgely reports seeing what he believes were pata- 
gonica at El Llano, eastern Province of Panama, on August 2, 1972, 
(6 birds, adults and immatures) and 2 on August 5 at Empire Range 


36 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


near Pedro Miguel Locks. N.G. Smith (Willis and Eisenmann, 1979, 
Smiths. Cont. Zool., no 291, p. 24) saw 4 at Barro Colorado Island, 
Canal Zone, on August 20, 1972, that were probably of this race. 


STELGIDOPTERYX RUFICOLLIS (Vieillot): Rough-winged Swallow, 
Golondrina Ala de Sierra 


FIGURE 3 


Hirundo ruficollis Vieillot, Nouv. Dict. Hist. Nat., nouv. ed., vol. 14, September 
1817, p. 523. (Bresil = vicinity of Rio de Janeiro.) 


Above, grayish to blackish brown, darker on remiges and rectrices 
than on back; throat light brown to grayish white, abdomen white to 
yellowish white. 

Description.—Length 125-132 mm. Adult (sexes alike), above gray- 
ish to blackish brown; throat variable in the subspecies from cinnamon- 
buff to grayish white; breast light brown, rest of undersurface white. 
Rump varying in subspecies from brownish to whitish. Adult male 
(and female, to less degree) with the tips of the barbs on the outer 
border of the first primary stiffened and recurved, so that they are 
roughened to touch. 

These are the most common and widely distributed of the swallows 
of Panama, ranging in open areas from the lowlands to the slopes of 
the mountains. In forested regions they occur along streams and 
around clearings. 

The whitish-rumped resident subspecies is seen often in pairs along 
roads, especially where these are bordered by cut banks in which there 
are shallow cavities that may shelter nest sites. In the period of north- 
ern winter the 2 dark-rumped migrant races from the north range in 
small flocks along the larger streams through the lowlands. 

The taxonomy of this complex remains in dispute, both as to the 
number of species and subspecies to be recognized. Here all forms are 
treated as conspecific. 

[Stiles (Auk, 1981, p. 282) has studied the systematics of these 
swallows in Costa Rica, concluding that 2 species are represented. Be- 
cause he cites Wetmore’s unpublished manuscript for the present vol- 
ume, we have left Wetmore’s account essentially unaltered. Basically, 
there are 2 distinct forms of Stelgidopteryx: a northern one that is uni- 
form brown above and mostly whitish below (serripennis group), and 
a more colorful southern one with a white rump, rufous throat, and 
yellowish underparts (ruficollis group). Stiles maintains that the 2 
groups behave as distinct species in Costa Rica, with serripennis being 


FAMILY HIRUNDINIDAE 37 


found in montane regions and ruficollis at lower elevations. There 
are localities where they evidently form mixed colonies without inter- 
grading, although it would seem that specimen evidence for lack of 
intergradation at these sites is scant. The problem remains with what 
to do with the form decolor, which has a rather circumscribed range, 
being found along the southern half of the Pacific coast of Costa 
Rica south and east in Panama to Veraguas. Stiles recognized decolor 
as a subspecies of Stelgidopteryx ruficollis, but its characters are 
exactly those one would expect of intergrades between ruficollis uro- 
pygialis and serripennis; such an interpretation is reinforced by the 
great variability observed in decolor. Based purely on museum speci- 
mens, one could argue that serripennis and ruficollis represent former 
isolates that in coming in contact along the Pacific Coast have formed 
a hybrid zone (there apparently are no pure parental types in the range 
of “decolor’’), whereas in their area of contact in the montane regions 
of the Atlantic slope they perhaps behave differently. More fieldwork 
and collecting will definitely be needed before this problem can be re- 
solved satisfactorily. S. L. O.] 


Ficure 3.—Portion of outer primary feather of the Rough-winged Swallow, 
Golondrina Ala de Sierra, Stelgidopteryx ruficollis, showing hooklike leading 
barbs. 


STELGIDOPTERYX RUFICOLLIS SERRIPENNIS (Audubon) 


Hirundo serripennis Audubon, Orn. Biogr., vol. 4, 1838, p. 593. (Charleston, South 
Carolina.) 


Characters —Throat grayish white; rump and upper tail coverts 
grayish brown concolor with back; pale edgings on secondaries re- 
duced in extent, and darker, less definitely white; upper surface some- 
what darker; wings dull black; tail brownish black. 

Measurements.—Males (10, taken in the breeding range, March to 


38 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


June), wing 105.2-111.4 (108.6), tail 47.0-50.6 (48.6), culmen from 
base 9.8-10.6 (10.2), tarsus 10.3-11.7 (11.2) mm. 

Females (10, taken in the breeding range, April to July 2), wing 99.7- 
104.3 (102.0), tail 45.3-49.5 (47.0), culmen from base 9.3-10.6 (10.2), 
tarsus 10.3-11.8 (11.2) mm. 

Winter migrant from the north, found locally along the lower chan- 
nels of the larger rivers. This is the breeding form of the eastern 
United States, southern Ontario, and southwestern Quebec. In Pan- 
ama, in the season of migration and the northern winter it is found 
regularly in small flocks along the larger stream channels in the low- 
lands. Museum specimens examined include 5 males and 12 females, 
collected from December 14 to March 8, in Bocas del Toro (Almirante, 
Changuinola), western Colon (Rio Indio), and northern Canal Zone 
(Juan Mina, above Gamboa, and Gattn). 

[I entertain doubts about separating the subspecies pulpit and 
psammochroa from serripennis but A. R. Phillips recognizes all three 
and has identified specimens of each from Panama. Many of those 
identified as psammochroa by Wetmore were considered to belong to 
serripennis by Phillips. We have not located an account or synonymy 
for the race fulvipennis among Wetmore’s manuscript material and it 
is not clear whether its omission was an oversight. In his card file is a 
hand-written entry (dated July 1958) indicating that he considered a 
specimen from Calovévora, Veraguas, in the British Museum to be re- 
ferable to fulvipennis, but in the Smithsonian collections the only speci- 
mens so labeled by Wetmore are a few very old ones from Costa Rica. 
An adult female specimen taken by Wetmore on June 10, 1953, at 
Llano del Jardino, 15 miles east of Sona, Veraguas, he originally identi- 
fied as uropygialis and then as a hybrid between uropygialis and fulvi- 
pennis. Wetmore was doubtless influenced by the mid-summer date, 
however. As the specimen shows no trace of any of the characters of 
uropygialis, it would have to be assigned to serripennis in the broad 
sense. It may represent a bird that simply failed to migrate north in 
the springs Ss: 1o @p| 


STELGIDOPTERYX RUFICOLLIS PSAMMOCHROA Griscom 


Stelgidopteryx ruficollis psammochrous Griscom, Proc. New England Zool. Club, 
vol. 11, December 14, 1929, p. 72. (Oposura, Sonora.) 


Characters.—Like Stelgidopteryx ruficollis serripenms, but averag- 
ing slightly paler on upper surface. Scapulars, upper wing coverts, and 
outer web of secondaries and inner primaries edged pale reddish brown. 


FAMILY HIRUNDINIDAE 39 


Measurements.—Males (10 taken in breeding season, March to May 
in southern Arizona), wing 104.5-111.5 (108.6), tail 45.3-50.0 (47.5), 
culmen from base 8.5-10.8 (9.7), tarsus 10.2-11.2 (10.6) mm. 

Iemales (10 taken in breeding season, April to June, in Baja Cali- 
fornia, southern California, and southern Arizona), wing 100.2-107.4 
(104.4), tail 42.9-48.7 (46.3), culmen from base 9.1-10.7 (9.9), tarsus 
10.2-11.5 (10.9) mm. 

Winter migrant from the north, found locally from December to 
March along the lower channels of the larger rivers. Recorded from 
specimens in western Colon (Rio Indio, Chilar) and the northern Canal 
Zone (above Gamboa, near Juan Mina). 

This is the form that breeds in the western United States and 
southwestern Canada. The series of 12 specimens taken in Panama 
from small flocks during January, February, and March in 1952, 1955, 
and 1961 includes 9 females and 3 males. 

In 1952, in February and March, I found these swallows abundant 
along the lower course of the Rio Indio in western Colon, in small 
flocks that fed over the water and the adjacent shore. In the main they 
ranged apart from the scattered pairs of the resident race uropygualis, 
which were more common farther inland along the river. 


STELGIDOPTERYX RUFICOLLIS UROPYGIALIS (Lawrence) 


Cotyle uropygialis Lawrence, Ibis, ser. 1, vol. 5, no. XVIII, April 1863, p. 181. 
(Atlantic slope, Canal Zone, Panama.) 

Stelgidopteryx ruficollis decolor Griscom, Proc. New England Zool. Club, vol. 11, 
Dec. 14, 1929, p. 69. (Divala, Chiriqui.) 


Characters.—In general like the northern migrant races, but throat 
and upper foreneck rufous; darker above; rump slightly to definitely 
paler than back, shading from light gray at base of feathers to nearly 
white at tips; breast, sides, flanks, axillars, and underwing coverts light 
grayish brown; carpal edge of wing barred rather indistinctly with dull 
white; center of lower breast yellowish white; abdomen and undertail 
coverts white, in some tipped with black. Immature, with back, wing 
coverts, and tertials edged (in part indistinctly) with dull cinnamon; 
rump, sides, and flanks tinged with cinnamon. 

In a female, taken at El] Llano, eastern Province of Panama, Febru- 
ary 3, 1962, the iris was wood brown; bill, tarsus, and toes black. In 
another of the same sex from Puerto Armuelles, Chiriqui, February 7, 
1966, the iris was dark brown, tarsus and toes fuscous, claws black. 

Measurements.—Males (10 from Canal Zone, eastern Province of 


4O BIRDS OF THE REPUBLIC OF PANAMA—PART A 


Panama, and Darién), wing 103.5-112.1 (108.0), tail 47.6-52.5 (50.3), 
culmen from base 9.1-11.0 (10.2), tarsus 9.6-10.9 (10.1) mm. 

Females (10 from Canal Zone, San Blas, eastern Province of Pan- 
ama, and Darién), wing 100.0-104.1 (101.0), tail 45.0-49.0 (47.3), cul- 
men from base 9.0-9.9 (9.3), tarsus 9.5-10.7 (10.1) mm. 

Resident. Ranges throughout, from the lowlands to 1500 m in the 
mountains of Chiriqui. They are seen commonly, singly, in pairs, or at 
the proper season, with groups of young, over streams or ponds, and 
also where earthen banks border roads. One or two may be noted 
among flocks of winter migrant swallows from the north, but in general 
these residents tend to feed and range apart. In late January 1957, at 
Mandinga, San Blas, 100 or more circled in loose company at sunset, 
feeding, and at intervals resting, in the open branches of a dead tree 
until early dusk, when suddenly they disappeared to sleeping quarters 
that I did not discover. At sunrise they came again to the tree to rest in 
the sun. When I collected 5 at random to check their identity on the 
morning of January 23, I was interested to find that all were females. 
Throughout the Isthmus these flocks are recorded through February. 
Then, in pairs or groups of half a dozen, they begin to examine the 
steep earthen banks along the rivers where later they nest. In the last 
week of March, along the Rio Jaqué in eastern Darién, they were start- 
ing to dig nest holes. Soon many were nesting, and through April their 
breeding season was fully under way throughout their range. In early 
June, near Sona, in the lowlands of western Veraguas, fully grown 
young were on the wing. 

Males at times were protective of their nesting area against intru- 
sion by other swallows. In late February 1960, on the Brackney farm 
at Palo Santo, Chiriqui, a pair was preparing to nest in a low bank on 
the shore of a small impoundment to water farm animals. The male 
dove constantly at the small Blue-and-white Swallows that came to 
drink and bathe. On one occasion he also pursued a single Vaux’s Swift 
that came down to the water. I was intrigued to observe that in the 
long, swinging circles of their flight, the swift, in escaping, did not out- 
distance the swallow. 

Like related swallows, these birds feed on small insects captured in 
endless circling in the air. The only call that I have heard from them is 
a low, harsh chip, a sound without great carrying power. 

I have noted that among this race of the Rough-winged Swallow 
some dig their nest holes while others may use old burrows made by the 
local kingfishers. The holes may be in the face of a perpendicular bank 
along a stream, or along a cut bank beside a roadway, away from water. 


FAMILY CORVIDAE Al 


The nest is a fairly compact mass of leaves, straws, and similar ma- 
terials, with a depression in the upper side for the eggs. These are white 
and may be from 4 to 6 in number. Most of the published observations 
of nesting in this swallow deal with the form or forms found in Guate- 
mala or in Trinidad. According to Belcher and Smooker (Ibis, 1937, 
pp. 506-507), the subspecies aequalis of Trinidad, similar in size and 
color pattern to uropygialis of Panama but with whiter rump, white 
edgings on the longer secondaries and inner primaries, lighter breast 
and yellow abdomen, has the eggs “immaculate white, smooth-shelled 
and slightly glossy.’ A set of 3 measured 18X13, 19X13, and 18.5x 
12.5 mm. The measurements cited for the race uropygialis in Schon- 
wetter, Handbuch der Oologie (Lief. 17, 1970, p. 194), of 18.5-20.3 x 
13.0-14.5 mm, average somewhat larger. 

The colors recorded in the description above are those typical of 
uropygialis in Panama from Darién and San Blas west on both Carib- 
bean and Pacific slopes through Veraguas; beyond, through Chiriqui 
and Bocas del Toro, and continuing in Costa Rica and Nicaragua to 
southeastern Honduras (Howell, Condor, 1972, p. 326). In addition, 
there is much individual variation toward the duller colored subspecies 
fulvipennis, found in typical form farther north in Central America. 
These individuals, with variable, generally paler coloration, were sepa- 
rated by Griscom as another race, decolor. They are found regularly 
through the western sector of the Panama range of the normally colored 
uropygialis and appear to have no valid racial status. 


Family CORVIDAE: Crows and Jays, Cuervos y Urracas 


The Corvidae, a family of more than 100 species, are distributed 
widely through all continents of the World, except Antarctica. It is 
interesting that the prominent crow-raven group, of large size and 
mainly black color, found through much of the Old World, even to 
Australia, in the Americas does not extend south beyond northwestern 
Nicaragua. The smaller jays range into South America as far as Bo- 
livia. 

The jays are omnivorous and feed adeptly in a variety of fashions, 
even pursuing aerial insects in the manner of a flycatcher. They are 
usually found in small flocks that move noisily through the woods, 
traveling at all heights. Nothing seems to be recorded about the nesting 
behavior of any species in Panama, although such research would prove 
most rewarding. Not only are the varying degrees of nesting coopera- 


42 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


tion in tropical jays of interest, but even within a single species, such 
as the Brown Jay, the level of participation by morphologically im- 
mature birds seems to differ by region. 


KEY TO SPECIES OF CORVIDAE 


1. Smaller, wing less than 130 mm; wings and tail extensively blue......... Z 
Larger, wing more than 160 mm; wings and tail blacker, with little or no tinge 
Of “DM eer ie ace de See sae Hea Ane thas net a 3 
2. Throat and foreneck dull black; crown and back of head light blue; throat 
black. Azure-hooded Jay, Cyanolyca cucullata cucullata. p. 50 
Lower throat, foreneck, and a transverse band between the eyes, extending 
back narrowly on the side of the crown, light blue. 
Silvery-throated Jay, Cyanolyca argentigula argentigula. p. 48 
3. Foreneck and upper breast deep black; lower breast and abdomen pale yellow; 
a line on side of jaw and spots above and below eye light blue. 
Black-chested Jay, Cyanocorax affinis zeledont. p. 45 
Foreneck and upper breast dull brownish black, shading to dull white to light 
brown on rest of lower surface. 
Brown Jay, Psilorhinus morio cyanogenys. p. 42 


PSILORHINUS MORIO CYANOGEWNYS Sharpe. Brown Jay, Pia-pia 


Psilorhinus cyanogenys Sharpe, Cat. Birds Brit. Mus., vol. 3, 1877, p. 140, pl. 9. 
(Pearl-Bay Lagoon, Mosquito = Laguna de Perlas, Caribbean coast of Nica- 
ragua. ) 

Psilorhinus mexicanus captus Kennard and Peters, Proc. New England Zool. 
Club, vol. 10, August 25, 1927, p. 2. (Chiriquicito, Bocas del Toro, Panama.) 


Large; generally dull in color, with long-tipped tail; bill black, or in 
immatures, yellow; short bristly frontal feathers above base of bill 
ChECE: 

Description.—Adult (sexes alike), head, neck, and upper breast dark 
sooty brown, becoming dark grayish brown on back and scapulars, in 
some, paler on the upper tail coverts; wings and tail deep olive-gray, 
the inner rectrices usually darker; tail tipped rather broadly with white; 
lower breast changing to grayish brown, in some the shift in color 
gradual, in others abrupt; undertail coverts grayish white, occasionally 
nearly white; underwing coverts and scapulars light gray to dull white. 

There is a membranous pouch on the upper breast which, when the 
bird calls, is inflated so that it is briefly visible. 

Measurements.—Males (9 from Bocas del Toro), wing 178.0-200.0 
(184.1), tail 175.0-190.0 (187.9, average of 8), culmen from base 39.6- 
45.4 (42.8), tarsus 45.3-49.0 (47.5) mm. 

Females (7 from Bocas del Toro), wing 180.0-193.0 (188.4), tail 


FAMILY CORVIDAE 43 


172.0-198.0 (185.1, average of 6), culmen from base 40.6-42.7 (41.6), 
tarsus 46.4-52.2 (48.5) mm. 

Resident. Fairly common locally through the lowlands of Bocas 
del Toro, from near the Rio Sixaola west to Chiriqui Grande. 

Brown Jays range usually in small bands, in the borders of open 
forest and second growth; they are not found in extensive stands of 
heavy forest. Commonly they move about under cover among leafy 
branches where they may not be noticed. They often fly out singly, 
however, or as a group across open areas, calling loudly and making a 
sharp popping sound through quick inflation and deflation of the curi- 
ous air sac exposed on the upper breast. Lawton (in Jansen, 1983, 
Costa Rican Natural History, pp. 573-574) reports that flocks use 8- 
to 10-ha home ranges, defending a territory within the home range 
only when breeding. The limited area where the species is known at 
present in Panama marks the southern extension of the species, which 
is found on the Caribbean slope of Central America, north to north- 
eastern Mexico in Nuevo Leon and to extreme south Texas. It crosses 
to the Pacific slope only in northwestern Costa Rica. 

Adult individuals have the bill and feet black. In immature birds 
these areas are yellow to partially black, the pattern varying apparently 
according to age. 

Lawton (in Jansen, 1983, Costa Rican Natural History, pp. 573-574) 
found that in Costa Rica young members of the flock help build the 
nest and will occasionally sit in it, but their efforts are irregular. The 
number of females that contribute eggs to the nest or take part in incu- 
bation varies. In one case where more than one bird was known to have 
contributed eggs to the nest, only one bird was seen to incubate or 
brood. The average clutch size of 30 nests examined by Lawton was 
4.5, nearly twice that reported by Skutch (Pacific Coast Avif., no. 30, 
1960, pp. 231-257), further suggesting wide variability in the breeding 
habits of this species. 

In Guatemala, Skutch recorded a single brood each year, beginning 
in February and March. The nest site was in trees in pastures or in 
recent second growth, with the nests high and inaccessible near the tips 
of long, slender branches. Other locations were in high forks in trees, 
and in the crowns of banana plants. The bulky nest is made of sticks, 
with a shallow cup lined with long, fibrous roots pulled by the birds 
from the earth. Construction, by both male and female, is accompanied 
by much calling. Lawton and Lawton (Auk, 1980, pp. 631-633) have 
shown that critical factors in nest-site selection are protection from 
wind and isolation from predators. Thus, the birds seek sites relatively 


44 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


sheltered from wind, but also isolated from other trees that would give 
predators easier access. Lawton and Lawton found that nests in more 
wind-protected sites were more likely to be successful than nests in 
more exposed sites. 

The eggs are bluish gray “thickly covered with fine brown speckles, 
which on the thicker end nearly obscure the ground color.” Seven eggs 
measured from 32.5-36.5X23.8-25.4 mm. The incubation period 
ranged from 18 to 20 days. The young at hatching have a yellow skin, 
sometimes darker on the upper surface, and are without down. Bill and 
feet are yellow, and the inside of the mouth is red. Skutch noted that 
the bills of the immature birds darkened slowly with age. 

Lawton and Guindon (Condor, 1981, pp. 27-33) found that all mem- 
bers of the flock bring food to the nest. Over the course of the nesting 
season inexperienced young birds become more efficient at bringing ap- 
propriate food items. 

The curious inflatable breast sac found in all races of this species of 
jay appears to have been noted first by Samuel Cabot, Jr. (Journ. Bost. 
Soc. Nat. Hist., vol. 4, 1844, pp. 460-467), in the race P.m. vociferus in 
Yucatan. He speaks of their calls as loud and disagreeable and in a 
male and female that he collected, records “‘a most peculiar formation 
in the trachea, being a membranous bag, coming off between the rings, 
about halfway down, and intimately connected with the skin of the 
neck.” There can be no doubt that he noted the peculiar air sac, though 
his interpretation of it as connected with the trachea was in error. 

Lee S. Crandall (Zoologica, vol. 1, no. 18, 1914, p. 337) in observa- 
tions near Guapiles, on the base of Volcan Turrialba, Costa Rica, in 
early 1914, made careful observations on the air sac in the field, verified 
later by data from a captive bird at the New York Zoological Society’s 
Bronx Zoo. He described the popping noise heard in the field, and in 
the zoo recorded that the bird “never uttered a vocal note, but distended 
the cervical sac whenever he was excited, making a popping sound which 
could be heard at a distance of several yards. On examination of this 
bird after death, the sac was found lying between the branches of the 
furculum, 1 mm anterior to their point of union. Deflated, it measured 
13 mm from base to tip and 19.5 mm along the base, the tip being 
rounded. ... The sac communicated directly with the praebronchial or 
interclavicular air-sac (Saccus interclavicularis ), through a large open- 
ing in the furcular membrane, and doubtless received its air from this 
source.” 

A more detailed account is that of Sutton and Gilbert (Condor, 
1942, pp. 160-165, figs. 59, 60) based on observations and specimens 


FAMILY CORVIDAE A5 


from northeastern Mexico. Their illustration of a plucked specimen 
shows clearly the position of the distended sac, which externally pro- 
jects from the space between the bifurcation of the two arms of the 
furcula. This verifies the anatomical details given by Crandall, with the 
appropriate suggestion that to avoid confusion with terms current for 
the air sac system in birds it be called the furcular sac. 

During fieldwork in southern Veracruz, Mexico, in early 1939, I 
found the nominate race, Psilorhinus m. morio, common, and heard 
their curious explosive sounds regularly. When they flew overhead, a 
yellowish spot appeared on the breast, though at rest this was not visible. 
With birds in the hand I found that the furcular sac was bare except 
for a few scattered filamentous feathers, mainly on the lower portion. 
The skin of the upper, anterior half was thin and partly transparent. 
The thicker-walled lower end, measuring approximately 15x20 mm, 
was dull buff in color. Posteriorly, the sac narrowed immediately to a 
slender tube as it entered the breast behind the furcula, and continued 
thus to its connection with the interclavicular air sac. In jays recently 
dead, I could inflate and deflate the sac by compressing and releasing 
the posterior part of the body so that air was forced into it and then 
withdrawn (Wetmore, Proc. U. S. Nat. Mus., vol. 93, 1943, pp. 298- 
299). 

Hardy, in his revision of the New World jays (Condor, 1969, p. 
363) lists the Brown Jay in the genus Cyanocorax, subgenus Psilor- 
hinus, as Cyanocorax morio. As the curious furcular sac has no known 
approach in any other species of the family, it seems appropriate to 
place this jay in a distinct genus as Psilorhinus morio. 


CYANOCORAX AFFINIS ZELEDONI Ridgway: Black-chested Jay, 
Chocho 


FIGURE 4 


Cyanocorax affinis zeledom Ridgway, Auk, vol. 16, no. 3, July 1889, p. 255. (Tala- 
manca, Costa Rica.) 


Fairly large; head, foreneck, and upper breast black; rest of lower 
surface pale yellow; wings and tail bluish; tail with a broad white tip. 

Description.—Adult (sexes alike), head, foreneck, and upper chest 
black; spot on posterior half of superciliary area, another on posterior 
half of lower eyelid, and a malar streak, bright blue; nape purplish 
blue; lower hindneck, back, scapulars, and rump dull, somewhat bluish, 
brown; wings, upper tail coverts, and tail dull blue, the tail tipped 
broadly with pale yellowish white. 


46 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Juvenile, crown and side of head dull blackish brown; throat, fore- 
neck, upper breast, hindneck, scapulars, and back dull grayish brown. 

An adult male taken at Las Palmitas, Los Santos, January 24, 1962, 
had the eye yellow; bill, tarsus, and toes black. In a fully grown im- 
mature male shot on the same day, the iris was duller yellow, and the 
external edge of the gape honey yellow. Other markings were as in the 
adult. 

Measurements.—Males (10 from Chiriqui to Darién), wing 163.0- 
175.0 (167.5), tail 157.0-168.0 (162.1), culmen from base 30.3-36.2 
(33.1), tarsus 45.0-52.0 (47.8) mm. 

Females (10 from Chiriqui to Darién), wing 159.0-169.0 (164.3), 
tail 154.0-161.0 (158.3), culmen from base 31.0-35.9 (33.7), tarsus 
47.0-51.9 (50.1) mm. 


Figure 4.—Black-chested Jay, Chocho, Cyanocorax affinis zeledont. 


Resident. Common in forested areas (including second growth) 
throughout the Republic, from near the Costa Rican boundary in Chiri- 
qui (Divala, Santa Clara) and Bocas del Toro (Rio Sixaola, Changui- 
nola) including the Azuero Peninsula (Pedasi, Tonosi, Las Palmitas) 
on both slopes to the Colombian boundary, to 1300 m elevation (near 
El Volcan and 1600 m (above Boquete at Horqueta, Velo); 600 m 
on Cerro Pirre (Cana) in Darién. 

As these jays range in forest and brushy cover, when these are 
cleared in cultivation, the birds disappear. However, if the clearings 


FAMILY CORVIDAE 47 


are later abandoned so that they are covered gradually by second growth 
(rastrojo), the birds may return. They seem to prefer edges and 
second growth to mature forest. The species is no longer found in 
Barro Colorado Island; this may be an indication that it avoids solid 
forest. 

Customarily they range in small groups, scattered through the forest, 
usually under cover. In early morning they may come out into border 
fields to rest in the sun in dead trees, but as the morning advances, dis- 
appear again into forest cover. Usually from 3 to 6 or 8 are found in 
company, though once near Mandinga in western San Blas I encoun- 
tered a flock of 30 ranging through the border of a mangrove swamp 
where the trees grew in shallow water. 

Usually they move quietly, hopping or flying a few feet at a time 
along the tree limbs. If not alarmed, in crossing rivers or other open 
areas they begin with quick strokes of the wings and then continue by 
gliding. They are adept at this, and may sail gracefully for 30 m or 
more before they lose momentum. 

The country name of chocho is given in imitation of their usual call. 
This name was in common use among the Choco Indians, but was varied 
occasionally to a harsher sound, geo geo, by some of the Cuna. Though 
the birds are secretive, they come readily to the usual squeaking sounds 
that naturalists use to attract birds. Eisenmann interprets one com- 
mon call as kyuck-kyuck, or kyuck alone. 

Once as the members of a flock called steadily, I approached quietly 
to find them resting or clinging in close company a meter above the 
ground, peering down at a small snake partly concealed in the low 
vegetation. More rarely, I heard low calls of jay jay when the birds 
were not alarmed. Also, I occasionally saw them mobbing small hawks, 
once, to my surprise, one of the predatory Collared Forest-Falcons. 
Ridgely (in litt.) has seen them mob a roosting Spectacled Owl. 

Some of the stomachs that I examined held parts of small lizards, 
others the remains of large ants, beetles, and seeds and other partly 
digested vegetable matter. Three birds taken by Burton (Bull. Brit. 
Orn. Club., 1975, p. 85) weighed 194, 203, and 222 g. 

In many days afield where these jays were common, the only ob- 
servation connected with nesting was near Yepe on the Rio Paya, 
Darién, in mid-March, when one flew across the open river carrying a 
straw, followed closely by a companion. Goodwin (Crows of the 
World, 1976, p. 306) describes the nest as being of “sticks and twigs, 
lined with fine pliable twigs, and probably also at times with tendrils 
and fibres.”’ A nest found near Rio Parancho, lower Atrato Valley, 


48 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Colombia, was in “a small isolated tree covered with creeping spiny 
palms.” Goodwin gives the clutch as 3 to 5; the eggs are “pale buff or 
brownish white, spotted and blotched with olive brown and with under- 
lying lilac or grey markings.” Two eggs collected by T. K. Salmon at 
Remedios, Antioquia, Colombia (no date) now in the British Museum 
collection measure 34.6 X 24.6 and 34.0 X 24.65 mm. In the Santa Marta 
region of Colombia, nests with eggs have been found from early April 
until mid-May, and Hardy found a pair with dependent but nearly 
full-grown young in Venezuela in July. 

The nominate race affinis found in Colombia and western Venezuela 
differs in being clear white on the lower surface and the tip of the tail. 
The difference between the two forms is readily seen when series of 
birds from the two areas are examined. Occasional birds from eastern 
Darién and San Blas in eastern Panama show some approach to this 
lighter condition, but the population of northern Choco, Colombia, is 
like that of Panama and is assigned to zeledoni. 


CYANOLYCA ARGENTIGULA ARGENTIGULA (Lawrence): 
Silvery-throated Jay, Urraca Garganta de Plata 


Ficure 5 


Cyanocitta argentigula Lawrence, Ann. Lyc. Nat. Hist. New York, vol. 11, nos. 
3-4, February 1875, p. 88. (Near Pico Blanco, southern Cordillera de Talamanca, 
Costa Rica.) 

Cyanolyca blandita Bangs, Proc. Biol. Soc. Washington, vol. 19, July 30, 1906, p. 
109. (Volcan de Chiriqui, above Boquete, 2750 meters.) 


Description.—Length 255-265 mm. Adult (sexes alike), head, back, 
chin, sides of neck, breast, and undersurface of wings black; lower 
back, scapulars, rump, upper and undertail coverts, abdomen, and flanks 
dusky, with a dull purplish blue tinge; wings and tail dull blue; fore- 
neck broadly pale bluish white, with lower margin where it joins the 
upper breast paler, nearly white; a narrow transverse band of bluish 
white across the center of the crown at level of the eyes, extending 
laterally as a narrow line along the side above and behind the eyes on 
either side. 

A male and female, taken along the Boquete trail at 2100 m above 
Bajo Grande, beyond Cerro Punta, Chiriqui, March 4, 1955, had the 
bill black; inside of mouth dusky neutral gray; tongue neutral gray; 
iris dark brown; tarsus, toes, and claws dull black. 

Measurements.—Males (10 from Chiriqui), wing 117.8-129.5 
(122.7), tail 118.6-133.1 (126.6), culmen from base 27.4-30.8 (29.0), 
tarsus 34.5-37.2 (35.8) mm. 


FAMILY CORVIDAE 49 


Females (5 from Chiriqui), wing 118.5-121.1 (119.6), tail 118.7- 
128.0 (123.2), culmen from base 28.0-30.4 (29.0), tarsus 32.3-36.8 
(o0.1)) mm. 

Resident. Found locally on the upper slopes of Volcan de Chiriqui, 
from 1525 to 3000 m elevation, from above Boquete to above Cerro 
Punta. 


Ficure 5.—Silvery-throated Jay, Urraca Garganta de Plata, Cyanolyca argenti- 
gula argentigula, 


On the forested slopes of the higher levels of Bajo Grande, beyond 
Cerro Punta, I found this jay ranging through the higher branches of 
the forest trees in scattered flocks of a dozen to 20 or more. In man- 
nerism they suggested common species of the north of the genus Cyano- 
citta. Their low calls, pay pay pay, uttered slowly, also were similar in 
tone but less vociferous. While they did not seem timid, the fairly thick 
growths of the abundant epiphytes and subdued coloration of the birds 
tended to conceal their movements so that they were not conspicuous. 
When I took one or two for specimens, their companions seemed little 
alarmed. If their low calls were heard at a distance, the birds usually 
came near when I made the usual squeaking sounds of our bird calls. 
But against the dark background of the tree limbs and thick parasitic 
growths the birds often were so screened that they could not be seen. 
Their flights were for only short distances among the branches. The 
stomachs of those examined were filled with insects. 

The type specimen of the nominate race, argentigula, in the Smith- 
sonian collections, obtained in April 1874 by Juan Cooper, came from 
William M. Gabb. It appears to be the only older record known for 
this form for Costa Rica. Pitelka’s (Journ. Washington Acad. Sci., 


50 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


vol. 41, 1951, p. 113) thorough study of records of Gabb’s itinerary 
concluded that the specimen was collected “near and more or less north 
of Pico Blanco,” which is near the southern end of the Cordillera de 
Talamanca. Sipurio, which Pitelka mentions also, is lower down on the 
Caribbean slope in the area drained by tributaries of the Rio Sixaola. 
In older collections in the British Museum there are 2 specimens col- 
lected by Arcé, without other data, except that they are marked “Vera- 
gua,’ a term applied earlier to the whole of western Panama. 

Modern records with definite locality for argentigula, mainly those 
in the Monniche collection (Blake, Fieldiana: Zool., vol. 36, no. 5, 
1958, pp. 545-546), all lie on the Pacific slope. Bangs (Proc. New Eng- 
land Zool. Club, vol. 3, 1902, p. 57) listed a series of 11 taken by W. W. 
Brown in June 1901 from Volcan de Chiriqui, 3000 m. Bangs Joc. cit., 
p. 16) says that Brown collected “for a short time in June on the north- 
ern or Caribbean exposure of the volcano” from which the species has 
been included by Blake as from the Caribbean drainage. This is not 
certain, however, as Brown worked mainly near or above Boquete. 
From there the old trail to the Cerro Punta area was a usual route, 
which would have taken him into the area where I found this jay com- 
mon, on the Pacific slope. 

Eisenmann and others have seen this species frequently above Cerro 
Punta, western Chiriqui (on the trail to Boquete) between 1920 and 
2190 m, in pairs or small flocks, uttering a rather nasal, but not loud, 
chaak, cheuk, check. 

The race of the mountains of the Cordillera Central, farther north in 
Costa Rica, marked by paler, whiter throat and crown markings, was 
named Cyanolyca argentigula albior by Pitelka (cit. supra, p. 114) from 
a male in the Smithsonian collections obtained by Ridgway and Zele- 
don on Volcan Turrialba. 


CYANOLYCA CUCULLATA CUCULLATA (Ridgway): Azure-hooded Jay, 
Urraca Corona Azul 


Cyanocorax cucullatus Ridgway, Proc. U.S. Nat. Mus., vol. 8, May 6, 1885, p. 23. 
(Navarro, Costa Rica.) 


Crown light blue; rest of head and breast black, body dull blue. 

Description.—Forehead, side of head, upper breast, throat, hind- 
neck, upper back, and undersurface of wings black; crown and occiput 
light blue; back, wings, rest of lower surface, and tail dull blue. 

Iris maroon (or bright red, as in a specimen collected by Ridgely in 
Chiriqui); tarsus black (Kennard and Peters, Proc. Boston Soc. Nat. 
Hist., vol. 38, 1928, p. 465); bill black (note on specimen label). 


FAMILY CORVIDAE oe! 


Measurements.—Males (8 from Panama and Costa Rica), wing 
117.0-128.3 (121.6), tail 127.1-145.9 (136.6), culmen from base 24.7- 
33.0 (29.3 average of 7), tarsus 35.7-38.2 (37.0) mm. 

Females (8 from Panama and Costa Rica), wing 115.0-131.0 
(120.8), tail 115.5-140.3 (120.8), culmen from base 26.3-31.4 (29.1), 
tarsus 35.4-38.7 (36.6) mm. 

Resident. Known definitely in Panama from a male and female col- 
lected March 20, 1926, by Kennard at 900 m on the Boquete trail, in- 
land from Laguna de Chiriqui, and from 3 males recorded by Bangs 
(Proc. New Engl. Zool. Club, vol. 3, 1902, pp. 57-58) taken on the 
Caribbean slope at 2100 m by Brown in June 1901. Blake, in his ac- 
count of the Monniche collection (Fieldiana: Zool., vol. 36, 1958, pp. 
544-545) lists 2, male and female, taken June 26 and July 14 at Camp 
Holcomb on the Holcomb trail at 1500 m. 

The species is known in Panama definitely from Chiriqui and Bocas 
del Toro. The following early reports are of uncertain validity. Sharpe 
(Cat. Birds Brit. Mus., vol. 3, 1877, p. 127) under the heading Cyano- 
corax ornatus (an early name for C. cucullata) lists a specimen from 
“Veragua. M. EF. Arcé.” Apparently this is the basis for the report of 
“Panama. Veragua (Arcé)” by Salvin and Godman (Biol. Centr. 
Amer., Aves, vol. 1, 1887, p. 500, see errata and corrigenda, p. xliv). 
iideway (Bull. U.S: Nat. Mus., vol. 50, pt. 3, 1904, p. 323) in- 
cluded records of 2 specimens that need further consideration. One is 
a skin marked “Arcé, Veragua, 1877” (originally identified as Cyano- 
citta ornata) that came in the Boucard collection with no other informa- 
tion as to locality. The second is a male purchased from the mission- 
aries Heyde and Lux labeled “Chitra—Prov. Coclé, Isthmus of Pan- 
ama, June 18, 1889, irides red-brown, found higher parts of mountain 
virgin forest.” The detailed travels of these travelers are not clearly 
known, but it appears certain from available data that in the early part 
of 1889 they crossed from Nata, Coclé, to the Caribbean slope in the 
upper area drained by the Rio Coclé del Norte. Two of their principal 
localities were ‘“Cascajal” and “Chitra.” The region as yet is little 
known ornithologically. 

More recently, Ridgely (im litt.) found it fairly common in late 
February and early March 1976 at the Fortuna Dam site (900-1050 
m), east of Boquete, ranging the very humid foothill forest there in 
small groups and in pairs. The birds were rather noisy, but quite secre- 
tive and elusive, and only infrequently seen. He collected 1 on March 
4, 1976. 

N. G. Smith (in litt. to Eisenmann) reports seeing this species in 


52 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Bocas del Toro in April 1979 when he crossed the divide at 1700 meters 
from Cerro Colorado, eastern Chiriqui. 

This race also occurs on the Caribbean slope of Costa Rica; other 
forms are found north to eastern Mexico. 


Family CINCLIDAE: Dippers, Mirlos Acuaticos 


The 5 species of this family, all of small size with heavy bodies and 
short tails, in general appearance suggest wrens. They are peculiar in 
the Order Passeriformes for their specialization for aquatic life, espe- 
cially through their dense, water-repellant plumage. Dippers are found 
along swift streams in hill country, where they enter the water to range 
along the bottom in search of food. In this they move by using the 
wings as in flight, or walk through skillful use of oblique currents and 
eddies made by stones and similar obstacles. They spend their lives en- 
tirely along streams, even through colder seasons, unless ice prevents 
their entering the water. 

Their bulky, domed nests may be placed at the border of waterfalls, 
or sheltered by stones along swift-flowing channels. The glossy white 
eggs, rather large in size, range from 4 to 6 in number. From observa- 
tions, as yet somewhat limited, it appears that for a period during molt 
the primary and secondary wing feathers may be renewed in such a 
manner that the bird may be incapable of aerial flight, but is still capable 
of its usual aquatic activities. 

The 5 species known are widely distributed—one in the Andes of 
South America; another in southern South America; one in Central 
America and western North America; one in Europe, northwestern 
Africa, and central and southern Asia; and a fifth in eastern and south- 
ern Asia. 


CINCLUS MEXICANUS ARDESIACUS Salvin, American Dipper, 
Mirlo Acuatico Americano 


FIGURE 6 


Cinclus ardesiacus Salvin, Ibis, vol. 3, no. 1X, January 1867, p. 121, pl. 2. (Cordil- 
lera de Tolé, inland from Tolé, Chiriqui, Panama.) 


Medium size; heavy body, short tail; adult gray, immature with 
undersurface white to grayish white. 

Description.—Length 165-180 mm. Adult (sexes alike), crown and 
sides of head dull brownish slate, shading on hindneck to the slate gray 
of the upper surface, this including the scapulars; wings and tail 


FAMILY CINCLIDAE 53 


blackish slate, margined with dull slaty; carpal edge of wing white; 
underwing coverts slate gray; a small white spot on upper and lower 
eyelids; chin and throat pale brownish gray; rest of undersurface light 
gray, slightly darker on flanks and undertail coverts, the latter tipped 
lightly with white. 

Immature, above faintly lighter gray; crown grayer; primaries, sec- 
ondaries, and greater and middle wing coverts tipped lightly with gray- 
ish white; undersurface white, except undertail coverts, which are gray. 

Measurements.—Males (10 from Chiriqui and Costa Rica) wing 
84.7-88.0 (86.3), tail 42.9-46.9 (44.9), culmen from base 20.5-22.2 
(21.4, average of 8), tarsus 31.1-32.9 (32.0) mm. 


Figure 6.—American Dipper, Mirlo Acuatico Americano, Cinclus mexicanus 
ardesiacus. 


Females (10 from Chiriqui and Costa Rica), wing 78.8-82.1 (80.7), 
tail 37.0-44.8 (40.2), culmen from base 20.2-21.4 (20.8, average of 9), 
tarsus 30.2-31.6 (31.0) mm. 

Resident. Recorded in small numbers along mountain streams. On 
the Pacific slope it is found from above Santa Fé, Veraguas (Eisen- 
mann) and in Chiriqui, from the Cordillera de Tolé west to the Rio 


54 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


Caldera, near Boquete, and the upper Rio Chiriqui Viejo, beyond El 
Volcan, from 1200 to 1700 m (reported by Brown, but perhaps un- 
certainly, to 2200 m). | 

In July 1965, Dr. C. R. Schneider (a chemist) believes he saw a dip- 
per on the Cana River near Cana, Darién (over 900 m) (Ridgely 1976, 
p. 265). If any dipper does occur in Darién it may be either this species 
or C. leucocephalus of the Andes. 

The Panama race is better known in Costa Rica, where it is found on 
the Caribbean slope of the central highlands as well as on the Pacific 
side of the mountains. In Panama it has not yet been recorded on the 
Atlantic drainage. It was described originally from 2 females—one 
adult, the other in immature dress—sent by Arcé to Salvin, from the 
mountains inland from Tolé. The type locality, a short distance west of 
the boundary with the Province of Veraguas, is the most easterly con- 
firmed report for the dipper in Panama. The sex of the 2 specimens 
which I examined in the British Museum (Natural History) is marked 
as female on the original labels. 

Another specimen in the British Museum, dated 1873, received 
from Boucard with the locality indicated only as “Panama,” may have 
come from the same source. These were the only specimens known 
until Brown collected a series of 11, near Boquete, and above on the 
slopes of the volcano, between February and April, 1901 (Bangs, 
Proc. New England Zool. Club, 1902, p. 51). The Monniche collection 
in the Field Museum has 3 specimens, 2 from Bajo Mono, and 1 from 
the Rio Caldera, the localities being at about 1580 and 1650 m. In the 
Smithsonian collections there is an adult female taken near Cerro 
Punta, Chiriqui, March 9, 1962, by Dr. C. L. Hayward. I recorded 
dippers occasionally, singly or in pairs, along the upper Rio Chiriqui 
Viejo from near the bridge west of EF] Volcan to the upper courses be- 
low Cerro Punta. They were not observed on the narrower channels 
above the village. Always they were wary, ranging along swift water 
in rocky channels. No specimens were collected, since it would not have 
been possible to retrieve them from the fairly deep, swiftly flowing 
water. Their habits and low calls were like those of the slightly darker 
race of the mountain streams of western United States. 

In March 1955, Glen Lewis told me that a pair came regularly along 
the river canyon beside his house below Cerro Punta. They had a nest 
concealed between two stones on the bank opposite his house, where it 
was inaccessible as there was no crossing. It appeared to be bulky, 
made of mosslike material. Eisenmann and Ridgely (in htt.) report 
nesting in the same general locality in 1969, 1972, and 1979. 


FAMILY TROGLODYTIDAE 55 


M.A. Carriker, Jr. (Ann. Carnegie Mus., vol. VI, 1910, p. 752), in 
his account of this species in Costa Rica, says of it that he found a pair 
on a river “high in the mountains above Ujarras de Terraba, where the 
stream came tumbling and foaming down through a deep gorge. Their 
discovery was caused by first finding the nest as I was climbing up a 
precipitous wall of rock beside a waterfall. It was a rather flat, cup- 
shaped structure, built almost entirely of moss and placed on a narrow 
shelf of rock on the brink of the falls. It contained one partially incu- 
bated egg which was blown, but later misplaced and lost... . In color 
it was plain dull white, of the usual shape and size of the Water Ouzel 
of the United States.” 


Family TROGLODYTIDAE: Wrens, Cucaracheros 


Wrens are widespread in Panama, where they are found at all eleva- 
tions. Twenty-two species, all permanent residents, are known from 
the Republic. They favor thick undergrowth and the lower levels in 
forests, where they feed on insects and other arthropods. The House 
Wren is common around human settlements. 

The family has a wide vocal repertoire; some members are among 
the finest singers in Panama, and an increasing number are known to 
sing antiphonally. Song is now being used as a taxonomic character; 
it has proved useful in determining the relationships of some variable 
and widely dispersed Neotropical species, such as Microcerculus mar- 
ginatus. 

Most wrens build domed nests with an entrance at one side; a few 
species nest in cavities. When not actually breeding, individuals of cer- 
tain species roost communally, often in a “dormitory” nest built for 
this purpose, or in a natural cavity. 

[Although now usually placed in the Mimidae, new evidence indi- 
cates that the monotypic genus Donacobiws should be included in the 
Troglodytidae. The species was at one time referred to as the “Wren- 
Thrasher” and was considered to be a wrenlike thrasher or a thrasher- 
like wren (Fuertes, Bird-Lore, vol. 13, 1913, p. 342), but its wrenlike 
attributes have of late been overlooked. Donacobius is considerably 
larger than most wrens, but is scarcely, if any, larger than the larger 
species of Campylorhynchus, and immatures, as well as the Bolivian 
race, D. a. albovittatus, which have a white superciliary stripe, bear a 
striking resemblance to Campylorhynchus griseus. The males and fe- 
males of Donacobius duet, which is a wrenlike trait. Olson and D. W. 


56 


BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Steadman examined skeletons of Donacobius and found a number of 
wrenlike characters, although in some other respects the osteology ap- 
peared to be intermediate between wrens and mimids. Mary H. Clench 
(pers. comm.) has found that Donacobius has the distinctive and 
unique pteryolosis of the Troglodytidae. It appears that on the basis of 
its specialized characters, Donacobius 1s clearly referable to the Troglo- 
dytidae, and it remains to be determined whether its other characters 
are merely primitive within the 10-primaried oscines or are indeed indic- 
ative of a derivation of the Troglodytidae from the Mimidae. S. L. O.] 


10. 


Ua 


12. 


KEY TO SPECIES OF TROGLODYTIDAE 


.. Dhreat: solid: black... saga See Ae 2 


Throat brown, white, buffy or streaked............. 2.2 ae 3 


. Black extending to breast. 


Black-throated Wren, Thryothorus atrogularts. p. 94 
Black limited to throat; breast chestnut. 
Sooty-headed Wren, Thryothorus spadix. p. 95 


. \Dhroat. white or -bufhy. cos cctools ec ee 4 


‘Throat brown or black and white. ».23)25) 00.2.5.) 922. oe 15 


. Head and underparts pure white. 

White-headed Wren, Campylorhynchus albobrunneus brachypterus. p. 60 
Crown “dark Or sy. os Bai BE OO 5 
Grown ‘blaeks Mauve. os eid etic SU ee See ae 6 
Crown: browaien sos cin) 5 od Seas ek Seki oe 0 ee 7 


. Total length more than 200 mm. 


Black-capped Donacobius, Donacobius atricapillus, p. 57 
Total length less than’ 160° mime. 20... 2 nd oo. oe or Z 


. Crown entirely black or washed with brown, total length 92-105 mm, tail very 


short. 
White-breasted Wood-Wren, Henicorhina leucosticta. p. 110 
Crown entirely black, total length 126-156 mm; tail longer, of normal pro- 
portions. Bay Wren, Thryothorus nigricapillus. p. 85 


. Back streaked black and white. 


Sedge Wren, Cistothorus platensis lucidus. p. 66 
Back: unstreaked .).. 000 ee CaaS sb A Se 8 


. Undersurface partly or entirely barred black and white........:..0..:- 9 


Undersurface not barred... 6. 2)k os oasis gos eg ool te 10 
Undersurface with band of solid black, abdomen barred black and white. 
Black-bellied Wren, Thryothorus fasciatoventris. p. 96 
Side of head and undersurface entirely barred black and white. 
Riverside Wren, Thryothorus semibadius. p. 92 
Breast definitely gray; tail very short. 
Whistler Wren, Microcerculus marginatus luscima. p. 121 
Breast whitish or washed with light, brown... ..)....-~- +2.) 12 
Superciliary prominent, whitew. la... ile. os bee eee se ee 13 


FAMILY TROGLODYTIDAE ly 


MCT NIN TTTCIIS Litt GE Ti esha aplasia it) faut ene abit nds UN oleae Sa bs ly dill RS alo 15 
13. Smaller, total length 95-100 mm. 
Timberline Wren, Troglodytes brown browm. p. 108 
PeiOra ln eneth UZ Oa tala. taiw de Shin a seve ele bate cea ale esi elele 6 14 
14. Uppersurface bright rufous; side of head barred. 
Rufous-and-white Wren, Thryothorus rufalbus castanonotus. p. 78 
Uppersurface, crown especially, grayish brown; side of head, below postocular 
stripe, white. Plain Wren, Thryothorus modestus. p. 72 
15. Superciliary indistinct, total length 108-115 mm. 
House Wren, Troglodytes aedon. p. 101 
Superciliary indistinct, total length 120-135 mm. 
Buff-breasted Wren, Thryothorus leucotis. p. 68 
EMMONS TECO AMI TEONVAA 1 Oe Fo, 588 ail er c4h 3 tig arcdrs heb eal Bh eibita obsue ties cS aud! GRMN Siallet Bataan le « V7 


17. Throat and breast rich chestnut; rest of body dark brown. 
Song Wren, Cyphorhinus aradus. p. 117 
Throat pale cinnamon, fading to buff on breast. 
Mountain Wren, Troglodytes solstitialis. p. 105 
18. Back barred black and white. 
Band-backed Wren, Campylorhynchus zonatus costaricensis. p. 64 
IBDSLE SOIGE 1 RON Aa 4 MI nee RRS PEORIA ede TNO AV) ane aU 19 
19. Back bright chestnut; breast gray; tail very short. 
Gray-breasted Wood-Wren, Henicorhina leucophrys. p. 114 
Back duller brown, breast not gray; tail of normal proportions........ 20 
20. Breast streaked black and white. 
Stripe-breasted Wren, Thryothorus thoracicus. p. 81 
SSF BASE | TST SN EKG OR i CU A TER aE SL A Pn Ee 21 
21. Breast tawny-rufous. 
Rufous-breasted Wren, Thryothorus rutilus hyperythrus. p. 99 
Breast gray-brown. 
Stripe-throated Wren, Thryothorus leucopogon grisescens. p. 83 


DONACOBIUS ATRICAPILLUS BRACHYPTERUS Madarasz, 
Black-capped Donacobius, Paralauta de Agua 


FIGURE 7 


Donacobius brachypterus Madarasz, Om: Monatsb. vol. 21, Feb. 1913, p. 22. (Ara- 
taca, Magdalena, Colombia.) 


Medium size; slender body, buff on lower surface, dark above, with 
crown black; long white-tipped tail. 

Description.—Length 210-235 mm. Adult (sexes alike), crown, 
side of head and hindneck black; back and greater wing coverts some- 
what dull brown; rump and upper tail coverts brownish buff; wings 
dull black, edged with dull brown; a white spot across base of primaries; 
tail black, tipped broadly with white, except on the central pair; entire 


58 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


undersurface pale buff, somewhat whiter on the throat; lower sides and 
flanks barred narrowly with dull black; axillars and anterior under- 
wing coverts dusky. 

A female of the nominate subspecies that I collected on the Rio 
Paraguay in northern Paraguay, September 30, 1920, had the bill black 
except for an elongate line of dull bluish gray around the base of the 
gonys; iris yellow; tarsus and toes fuscous; concealed bare skin on 
side of neck orange yellow. 

Measurements.—Males, (6 from Darién), wing 80.0-84.7 (83.0), 
tail 96.2-100.9 (98.9), culmen from base 24.9-28.5 (26.2), tarsus 32.0- 
34.6 (33.4) mm. 


Figure 7.—Black-capped Donacobius, Paralauta de Agua, Donacobius atri- 
capillus brachypterus. 


Females (2 from Darién), wing 79.5-80.3 (79.9), tail 94.5-94.7 
(94.6), culmen from base (one only) 23.2, tarsus 30.8-32.0 (31.4) mm. 

Resident. Recorded in Panama from FE] Real, Darién (specimens 
in the American Museum of Natural History include 6 males and 1 
female collected by Anthony and Richardson, January 1 to 8, 1915; 
male and female taken December 13, 1928). 

This species ranges in suitable lowland habitat from eastern Panama, 
northwestern Colombia, and most of South America, chiefly east of the 


FAMILY TROGLODYTIDAE 59 


Andes, to northeastern Argentina. The Panama race is recorded from 
Darien and northern Colombia to the Santa Marta region. 

On April 1, 1967, Eisenmann and H. Loftin found a small colony 
of three or more pairs near El Real, Darién, in a dry Montrichardia 
marsh with scattered bushes. Pairs displayed on leafless bushes, 
perched a few inches apart. In one pair watched at close range, the 
upper individual gave a whirrr or chirrr and spread the tail slightly as 
it moved from side to side; the lower bird sitting on a different twig 
answered with a louder, more musical, semiwhistled kweéa, kweéa, 
kweéa, also moving its tail. Eisenmann and Loftin did not notice the 
bare yellow skin, although they watched this pair perform three times 
at close range, once using a different perch, but in each case with the 
louder voiced bird sitting lower. 

On April 10, 1968, P. Schwartz took Eisenmann to a swampy pond 
in northern Venezuela, near the coast, where there was a small colony 
in a region of damp second-growth woodland. The birds perched on 
bushes in or near the pond. Both members of a pair perched on the 
same bush and vocalized antiphonally while wagging partly spread tails 
and sometimes exposing the bare orange-yellow neck skin. The semi- 
whistled call was noted as wd-ee (or hod-ee), wo-ee, wo-ee and the 
harsh call as a wheezing or buzzy wiizee, wiizee,wtizee. The differences 
in syllabification made in the field a year apart may or may not indicate 
a difference in vocalizations heard from 2 subspecies. 

Donacobius inhabits dense growths of grasses and rushes grow- 
ing in open fresh-water swamps. While I have worked at E1 Real sev- 
eral times, I have not found it. The marshy areas bordering the lower 
Rio Pirre are suitable habitat. 

Fuertes (PBird-Lore, vol. 15, 1913, p. 342) described the song of male 
Donacobius as a loud, whistled note like that of our Cardinal, with the 
female accompanying him with a low grunting call. He described and 
figured in both sexes an inflatable sac of bright yellow skin on the side 
of the throat “which when the bird sings, puffs out to the size of a 
cherry.” 

Skutch (Condor, vol. 70, 1968, pp. 74-75) in Venezuela, described 
the nest as a deep cup built around tall grass stems. The eggs “ap- 
peared to be uniform light, reddish brown. Closer scrutiny revealed 
that they were mottled rather than uniformly colored, but so densely as 
to cover the whole surface with slightly varying shades of reddish 
brown.” The “three eggs measured 25.5 by 16.5, 24.7 by 16.8, and 
24.4 by 16.3 mm.” The young at hatching were without down. 

Because of its uncertain taxonomy (it differs from the Mimidae, 


60 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


among which it was formerly placed, by various morphological char- 
acters and in long incubation and nestling period), Skutch suggests 
replacing the inappropriate English name ‘“‘Mockingthrush” by Dona- 
cobius. 


CAMPYLORHYNCHUS ALBOBRUNNEWUS (Lawrence), White-headed 
Wren, Cucarachero Cabeciblanco 


FIGURE 8 


A iarge, long-tailed wren; head and undersurface white; above dark 
blackish brown. Juvenile birds may have indistinct lines of pale gray 
in the crown. 

Description.—Length 180-200 mm. Adult (sexes alike), head, hind- 
neck, and undersurface (except as noted) plain white; underwing 
coverts, tibiae and undertail coverts spotted with dusky (in varying 
amount in the two races recognized here); rest of upper surface sooty 
brown. 

Immature (in juvenile plumage), crown and side of head slightly to 
heavily lined with dull grayish brown; tibia and undertail coverts 
spotted in varying amount with dusky; tail in some tipped narrowly 
and irregularly with white; duller, more brownish black above, with 
the upper back lined and spotted lightly with white; flanks, abdomen, 
and undertail coverts faintly to definitely washed with pale buffy brown. 
(This pattern of markings is followed by a plumage closely similar to 
that of the adult.) 

In recent reviews these wrens of Panama and northwestern Colom- 
bia are listed as geographic races of the species turdinus, widely dis- 
tributed in northern South America. The assignment has come through 
a peculiar population of wrens found in Narino in far southwestern 
Colombia, described by Meyer de Schauensee as Campylorhamphus 
albobrunneus aenigmaticus (Notulae Naturae, no. 209, Acad. Nat. 
Sci., Philadelphia, April 16, 1948). In the original description it is 
suggested only that the new form might indicate “a hybrid between C. 
albobrunneus and C. turdinus, but this possibility seems doubtful as 
the latter . . . is unknown west of the Andes. .. . neither species has 
been taken... . in Narino.” J. Hatter (Veroff Zool, ‘staatesamunale 
Munchen II, 1967, pp. 143-144) suggests that it is more probable that 
aenigmaticus is the result of hybridization between C. a. harterti and 
C. zonatus brevirostris of northwestern Ecuador. Meyer de Schauen- 
see (The Species of Birds of South America, 1966, p. 401) mentions 
both hybrid possibilities. Until there is clearer understanding it is pref- 
erable to consider albobrunneus, with its subspecies, harterti, as a sepa- 


FAMILY TROGLODYTIDAE 61 


rate species from C. turdinus and its races, all of which differ in adult 
stage in variable patterns of dark markings on the head and undersur- 
face. 

A record by Salvin and Godman (Biol. Centr. Amer., Aves, vol. 1, 
1880, p. 63) under the name “albibrunneus” from “Veraguas (Arcé)” 
quoted by other authors, is in error, as the species is not known from 
that area. 


yn 


iN 
J IN 
4 jp 
“ LPS 
\* , 


Y my ah 
SAN Nt 
PTY yt 


SS 
SS 
Ss 
JI 
IS 


> 


i, 

fe, 

rg Ny hy 
“ 


Ficure 8.—White-headed Wren, Cucarachero Cabeciblanco, Campylorhynchus 
albobrunneus. 


This handsome wren is an arboreal forest inhabitant that ranges also 
in trees and thickets bordering open lands. It is gregarious, so that 
usually 3 or 4 are found in company, ranging from near the ground to 
the median level, often 16 or 20 m up, or occasionally higher, in the 
trees. Usually they move under cover among the leaves, or through 
masses of creepers, where their presence is made known by their croak- 
ing, grating calls, though the birds themselves remain hidden. At times 
the notes are joined in a sequence that may be considered a song. When 


62 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


alarmed near the ground they move into higher branches with much 
fluttering of wings, resorting to flight only for distances of a few feet. 

When not disturbed the little parties may appear in small open areas 
on dry ground beneath thickets or low trees, where they dust in loose 
soil. Often, as one fluttered thus, a companion stood over it, apparently 
pecking at small parasites that appeared among the feathers of the 
crown. They may also be seen creeping into small openings among the 
exposed roots of fallen trees, such activities accounting for the soiled 
appearance often noted in the white feather areas. Less often, a small 
flock of as many as half a dozen flies across open trails, or between ex- 
panses of open pasture, moving quickly with rapidly tilting flight. At 
rest in branches they sometimes crowd closely against their companions. 
Where not disturbed, they rest on open perches, uttering their croaking 
calls, with fluttering wings and rapidly vibrating tails. 

In February, in the nesting season, I have seen groups of 3 flying 
openly among the top branches of tall trees in pursuit of one another, 
often crowding together, apparently in a form of territorial display. 

The nest and eggs are not known. 


CAMPYLORHYNCUS ALBOBRUNNEUS ALBOBRUNNEUS 
(Lawrence) 


Heleodytes albo-brunneus Lawrence, Ibis, Ser. 1, vol. 4, January 1862, p. 10. (Near 
the line of the Panama Railroad, near the summit of the Atlantic slope.) 


Characters.—Tibia paler, varying from white to light grayish 
brown; undertail coverts white, lightly spotted or barred with dull 
brown; immature with crown less heavily marked; where streaking is _ 
present, the lines averaging narrower, less distinct, and more grayish 
brown. 

Measurements.—Males (10 from northern Coclé, Canal Zone, east- 
ern Colon and Darién), wing 80.2-87.2 (83.4), tail 76.0-80.5 (78.3), 
culmen from base 21.3-25.7 (23.6), tarsus 24.3-26.7 (25.4) mm. 

Females (10 from northern Coclé, eastern Colon, and Darién), wing 
72.2-81.1 (78.0), tail 71.3-78.1 (75.6), culmen from base 21.1-24.3 
(23.5), tarsus 24.3-26.3 (25.00) mm. 

Resident. On the Pacific side, in the central valleys of the Rio Bay- 
ano, Rio Mamoni, and of the Rio Tuira (El Real, mouth of the Rio 
Paya, and the old village site at the base of Cerro Tacarcuna); Cana; 
Jaqué; on the Caribbean slope from northern Coclé, and the Canal 
Zone to eastern Colon. Although formerly recorded widely from the 
Chagres Valley of the Canal Zone, in recent years it has been observed 


FAMILY TROGLODYTIDAE 63 


in the zone only in the very humid forested northwestern area along 
the Achiote Road. 

This species is locally fairly common, but comes to attention mainly 
from its calls, as usually the birds move about under cover of leaves. 

A female, taken by E. A. Goldman at 850 m near Cana, May 22, 1912, 
had the stomach filled with insect remains that included chrysomelid 
and tenebrionid beetles, orthoptera, and ants. Three males collected by 
piauch (Bull. Brit. Orn. Club., 1977, p. 64) in the Province of Pan- 
ama and Canal Zone weighed 32.7, 34.0, and 34.3 g. 

Eisenmann (im litt.) found a group ot 3 or 4 on the Achiote Road 
on March 23, 1967, in shady, canopied, damp second growth ranging 
in the foliated trees from 5 to 20 m and giving harsh notes like churk 
or kurk. An unseen bird uttered a harsh song ending with semimusical 
kaw notes. At Altos de Majé above Rio Bayano, eastern Province of 
Panama, Eisenmann found 3 wrens on the ground, dust bathing in the 
forest; while dust bathing they were at times in physical contact. 


CAMPYLORHYNCHUS ALBOBRUNNEUS HARTERTI (Berlepsch) 


Heleodytes harterti Berlepsch, Ornis., vol. XIV, February 1907, p. 347. (San José, 
Rio Dagua, Valle, Colombia.) 


Characters —Feathered area of the lower leg (the tibia) with the 
markings definitely darker; undertail coverts duller white, heavily 
spotted and barred with dull grayish brown; immature, with brownish 
gray lines and feather borders on the crown and side of head more 
abundant, so that this area is darker. 

Measurements.—Males (10 from eastern San Blas, and adjacent 
Colombia, in Choco and Antioquia), wing 81.5-88.2 (85.3), tail 75.2- 
62.1! (79.5), culmen from base 22.5-27.4 (24.8), tarsus 25.5-28.1 
(26.6) mm. 

Females (9 from eastern San Blas and adjacent Colombia, in Choco 
and Antioquia), wing 76.0-82.5 (78.7), tail 72.7-82.5 (77.6), culmen 
from base 21.9-24.4 (23.3), tarsus 23.7-27.0 (24.7) mm. 

Resident. On the Caribbean slope, in eastern San Blas (Puerto 
Obaldia). The first record of this darker form was by Griscom (Bull. 
Mus. Comp. Zool., vol. 72, 1932, p. 358) of a female collected by von 
Wedel at Puerto Obaldia (date not listed). In the Museum of the 
University of Cincinnati there are 2 others, male and female, taken by 
Wedel at this locality August 28, 1931. 

It is possible that this race is found along much of the coast of San 
Blas, though recorded at present only at the eastern end. It is common 


64 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


in Choco, in adjacent Colombia, on both Atlantic and Pacific sides. Near 
the Pacific coast of Panama at Jaqué, along the Rio Jaqué near the 
Colombian boundary, and to a lesser extent on Cerro Pirre, harterti 
intergrades with nominate albobrunneus, occasional specimens being 
closely similar in the markings on the lower leg. A specimen, not de- 
termined as to race, was taken in February 1971 at Cerro Quia (300m), 
Darien; it is now the Gorgas Memorial Collection. 


CAMPYLORHYNCHUS ZONATUS COSTARICENSIS Berlepsch; 
Band-backed Wren, Cucarachero Listado 


FIGURE 9 


Campylorhynchus zonatus costaricensis Berlepsch, Auk, vol. 5, no. 4, October 1888, 
p. 449. (Costa Rica.) 

Heleodytes zonatus panamensis Griscom, Amer. Mus. Nov. no. 280, September 
10, 1927, p. 12. (Santa Fé, Veraguas.) 


Fairly large; with long tail, heavily spotted breast, and upper sur- 
face black, banded with white. 

Description.—Length 160-170 mm. Adult (sexes alike), crown pale 
brownish gray, spotted with black; upper surface black; hindneck 
streaked widely with white; back, wings, and tail banded broadly with 
white, the lines varied with buff, especially on the rump and tail; chin 
to upper breast white, becoming tawny-ochraceous on sides, lower 
breast, abdomen, and undertail coverts; foreneck and breast heavily 
spotted with black, these spots continued sparingly on sides and under- 
tail coverts. 

Juvenile, crown dull, slightly brownish, with a broad white super- 
ciliary stripe, and a black line behind the eye; upper surface black; 
wings and tail dull black, spotted and barred strongly with pale brown, 
the markings paler to nearly white on the primaries; edge of wing 
white; throat and upper breast, dull white, shaded lightly with buff; 
lower breast, sides, underwing coverts and undertail coverts cinnamon- 
brown. 

Measurements.—Males (10 from Bocas del Toro and Veraguas), 
wing 68.0-72.2 (71.6), tail 67.1-71.4 (68.5), culmen from base 20.9- 
2525,( 23:2), tarsus 22.8-2597 (2328) mame 

Females (10 from Bocas del Toro and Costa Rica), wing 66.2-71.5 
(68.1), tail 62.5-67.3 (64.9), culmen from base 19.8-21.9 (20.7), tarsus 
20.7-23.6 (22.4) mm. 

Resident. Found locally in Bocas del Toro (Almirante, Cricamola, 
upper Rio Changuena, and Veraguas (Santa Fé). 


FAMILY TROGLODYTIDAE 65 


The bird watcher familiar with the desert areas of southwestern 
United States and Mexico should easily recognize this species as a 
“cactus wren” from its colors and plumage pattern, but may be un- 
certain when he sees the birds ranging in tangles of vines, tropical 
shrubbery, and low-growing palms in fairly dense, humid jungle. 
Often they are found in small bands of half a dozen that move about 


Figure 9.—Band-backed Wren, Cucarachero Listado, Campylorhynchus zonatus 
costaricensts. 


with the usual curious jumble of croaking, rattling calls, differing 
mainly from their northern cousins in the darker pattern of their mark- 
ings. 

Skutch (Pacific Coast Avifauna, no. 34, 1960, pp. 186-201) gives a 
detailed account of their activities in Costa Rica and elsewhere in the 


66 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


American tropics. It is common for several, usually a family group, to 
live in company in forest. They feed actively in company through the 
day, and at evening take shelter also in company in a covered nest, 
specially built for such use near the tips of slender branches in forest 
trees. In the breeding season other nests are constructed, perhaps by 
the male alone, or by both members of the pair, the structure being 
generally similar to that used for sleeping. The eggs, three to five in 
number, vary from plain white to a pattern of faint markings of brown, 
which may form a wreath about the larger end. Measurements range 
from 20.6-23.8 by 15.1-16.2 mm. 

When grown, the family may continue to sleep in company. The 
companionship continues when the parents prepare a second nesting, as 
the grown young of the first brood may assist in care and feeding the 
second family. 

In Panama this species has been known mainly in the Chiriqui La- 
goon area, ranging inland, as indicated by a female taken by Galindo, 
prepared by Hinds, to over 700 m above the upper Rio Changuena in 
Bocas del Toro. While in Panama mainly a bird of the Caribbean slope, 
it crosses the Divide in the humid forested area still found above and 
north of Santa Fé, Veraguas. The species occurs in humid country 
northward to Mexico, and, with an interrupted range, is credited to 
northern Colombia and northwestern Ecuador. 

Griscom described a race based on slightly smaller size in 5 male 
specimens collected by Benson, March 4 and 6, 1925, at 670 m eleva- 
tion, near Santa Fé, Veraguas. The measurements are as follows (aver- 
age in parentheses): wing 65.7-72.2 (69.7), tail 66.9-73 (70.0), cul- 
men from base 20.9-25.5 (22.6), tarsus 23.1-24.5 (23.6) mm. A 
slight difference is noted, but so small that it has been considered of 
doubtful significance and has not been recognized as sufficient basis to 
warrant a separate race (see Hellmayr, Cat. Birds Amer., pt. 7, 1934, 
p. 140). 


CISTOTHORUS PLATENSIS LUCIDUS Ridgway: Sedge Wren, 
Saltapared de Ciénaga 


Cistothorus polyglottus lucidus Ridgway, Proc. Biol. Soc. Washington, vol. 16, 
November 30, 1903, p. 169. (Boquete, Chiriqui.) 


Very small; streak in front of eye; throat, foreneck, and breast 
mainly white; back, wings, and tail light brown with dull black barring. 
Description.—Length 95-105 mm. Adult male, crown and hindneck 


FAMILY TROGLODYTIDAE 67 


dark hair brown, streaked narrowly, especially on the crown, with 
whitish, the markings darker laterally, changing to blackish; back dull 
black, streaked with white; scapulars, lower back and upper tail coverts 
light brown, barred with black; rump more rufescent; tail dull black, 
tipped indistinctly with brownish white, barred on outer webs with 
brown; wings dusky, with outer webs barred with light brown, except 
at tips; sides of head brownish white, streaked lightly and narrowly 
with dull black; a narrow superciliary line dull white; throat, upper 
foreneck, lower breast and upper abdomen white; sides, flanks, tibia, 
and undertail coverts light brown; underwing coverts white. 

Adult female, back duller black; light markings of crown and back 
reduced in extent. 

Immature, above browner without pale markings on upper surface, 
or, if present, much reduced. 

Iris yellow; bill brown to blackish, sometimes yellow or whitish be- 
low; tarsi yellow to whitish or brown (labels on 7 Chiriqui specimens 
in the American Museum of Natural History). 

Measurements.—Males (10 from Chiriqui and Costa Rica), wing 
43.2-50.0 (45.7), tail 36.7-46.7 (45.7, average of 8), culmen from base 
W2:0-13.0 (12.5), tarsus 16.6-18.7 (17.6) mm. 

Females (10 from Chiriqui and Costa Rica), wing 43.5-46.0 (44.6), 
tail 36.4-45.0 (39.5), culmen from base 11.3-13.6 (12.8), tarsus 17.3- 
19.4 (17.9) mm. 

Resident. Recorded in small marshy savannas near Boquete and 
Bugaba, western Chiriqui, but judging by habitat in Costa Rica, may 
also inhabit dry grassy fields near montane ponds. 

The species is known as yet in Panama only through a few specimens. 
One, in the Smithsonian, received from Salvin, collected by Enrique 
Arcé, is marked only as from “Veraguas,”’ which at that time may 
have included Chiriqui. Salvin (Proc. Zool. Soc. London, 1870, p. 182) 
mentions one from Bugaba (a lowland locality), Chiriqui, collected by 
Arcé. Salvin and Godman (Biol. Centr. Amer. Aves, vol. 1, 1888, pp. 
106-107) wrote that of “Chiriqui examples we have now seen several.” 
In the bird collections in the British Museum there are 4, labeled ‘Chiri- 
qui’ without other data. A male in the Museum of Comparative 
Zoology was taken by W. W. Brown 1500 m above Boquete, April 25, 
1901. 

The American Museum of Natural History has 8 specimens from 
western Chiriqui, almost all collected in 1904 and 1905 by H. J. Watson. 
Six are labeled Boquete, at 2000, 3500, and 4000 ft., December 6, 21, 


68 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


1904, and March 7 and October 5, 15, 1905. Two are labeled “Volcan 
Chiriqut” 9000 ft. October 14, 16, 1904. 

This species has not been reported from Panama since the specimens 
mentioned above were collected. It may well be endangered. 

In Costa Rica the bird has been collected frequently, but with little 
information concerning its habits. Carriker (Ann. Carnegie Mus., vol. 
6, 1919 p. 772) noted that it was “found in only a few places in Costa 
Rica, but is abundant wherever it occurs. It makes its home in the 
grassy marshes which are found in the vicinity of La Estrella and 
Azahar de Cartago at an altitude of about 5000 feet. It has all the 
habits of the Marsh Wrens of North America.” 

The eggs of related races of the species platensis are described as 
white without markings (Meise, W., in Schonwetter, Handb. Ool., pt. 
19, 1971, pp. 330-331, plate 4, fig. 8). 

Slud (Bull. Amer. Museum Nat. Hist., 1964, no. 128, p. 282) de- 
scribes the call of this race in Costa Rica as a grasshopperlike tgrr and 
the song as a chyip-chyip-chyip-chyip-chyip, sometimes with a gurgling 
quality. 

It may well be that the complex of populations included in the species 
platensis should be divided into two or more species. Some of the 
South American forms are, by voice at least, very distinct. Tropical 
populations of this species are often known as the Grass Wren. 


THRYOTHORUS LEUCOTIS Lafresnaye: Buff-breasted Wren, 
Cucarachero Pechi-Amarillejo 


Thryothorus leucotis Lafresnaye, Rev. Zool., vol. 8, September 1845, p. 338. (“Co- 
lombia aut Mexico” = Honda, Rio Magdalena, Colombia, designated by Hell- 
mayr, Cat. Birds Amer., pt. VII, 1934, p. 165.) 


Rather small; upper surface definitely brown, brighter on rump and 
upper tail coverts; wings and tail russet, heavily barred with black; 
below ochraceous-buff to tawny-brown, darker on sides and undertail 
coverts; throat white. Similar in general to Thryothorous modestus but 
distinguished by heavy, much more distinct black bars on wings, and 
usually more contrast between white throat and buffy breast. 

Description.—Length 120-135 mm. Adult (sexes alike), crown 
dull brown changing to a deeper shade on back and scapulars, and be- 
coming brighter on rump and upper tail coverts; wings, including co- 
verts, russet brown, barred narrowly but distinctly with black; tail rus- 
set, barred heavily with black; a white line over eye from base of bill 
back along side of crown; a dull brown line back of eye; side of head 


FAMILY TROGLODYTIDAE 69 


below this, and the upper throat, white, with cheeks in some narrowly 
streaked or spotted with gray; white of throat changing to ochraceous- 
buff on foreneck and breast, deepening on sides, flanks, and undertail 
coverts to tawny brown; edge of wing, scapulars, and underwing co- 
verts white. 

Immature, colors somewhat duller, with the bars on wings and tail 
less distinct. 

In the Canal Zone and adjacent parts of the Provinces of Panama 
and Colon the Buff-breasted Wren may be found in the same wood- 
lands as the paler-colored Plain Wren, Thryothorus modestus, but 
avoids drier areas, being usually found near streams, or swampy or 
damp woods. It is present eastward on both slopes through Darién and 
San Blas to Colombia, with a local race in the Perlas group on Isla del 
Rey and adjacent islands. Beyond the Isthmus of Panama, in tropical 
South America, the species ranges widely with several races in the low- 
lands of Colombia south to Brazil and Peru. 


THRYOTHORUS LEUCOTIS GALBRAITHII Lawrence 


Thryothorus Galbraith Lawrence, Ann. Lyc. Nat. Hist. New York, vol. 7, June 
1861, p. 320. (Atlantic side on line of R.R.= Lion Hill, Panama Railroad, 
Canal Zone.) 


Characters.—-Dull brown dorsal surface, with tone more olive than 
rufescent, as is T. 1. conditus. 

Male and female, taken at La Jagua, eastern Province of Panama, 
March 22, 1961, had the iris wood brown; maxilla fuscous-black, ex- 
cept on the cutting edge, which, with the mandible, was pale dull neutral 
gray; tarsus and toes dusky neutral gray. 

Measurements.—Males (10 from Canal Zone, eastern Province of 
Panama, San Blas, and Darién), wing 62.0-65.4 (63.6), tail 41.0-44.3 
(42.8), culmen from base 18.5-20.6 (19.7), tarsus 23.2-24.9 (24.0) 
mm. 

Females (10 from eastern Province of Panama, San Blas, and 
Darien), wing 58.0-60.9 (59.6), tail 37.6-41.1 (39.3), culmen from 
base 17.0-19.6 (18.0), tarsus 22.3-24.3 (23.5) mm. 

Resident. Locally common from the Canal Zone east on the Pacific 
side through the eastern Province of Panama to Darién (at mouth of 
Rio Tuquesa, on the Rio Chucunaque, and at El Real and Marraganti 
on the Rio Tuira; Cerro Sapo); on the Caribbean slope from the north- 
ern Canal Zone east through eastern Colon and San Blas (Mandinga, 


70 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Permé). Not definitely recorded west of the Canal Zone, although 
present on the Achiote Road. No longer present on Barro Colorado 
Island (Willis and Eisenmann, Smiths. Cont. Zool., no. 291, 1979, 
p. 24). 

These wrens range usually in pairs in thickets, masses of vines, and 
other low growth, commonly near water. The song, heard regularly, is 
a loud, clear repetition of a phrase that may vary slightly. Occasionally 
two birds may join, with the calls of one following quickly after the 
other. Often one may answer another, even at some distance, their 
notes suggesting those of Thryothorus nigricollis but higher in pitch, 
in a repetition of three notes that vary in cadence in different indi- 
viduals. Their songs at times may suggest those of the Bay Wren. 
Frequently I have heard them scolding with low calls, chit-it, chip wit. 
Songs noted by Eisenmann in Panama City and in Darién were loud, 
ringing, and rich, very different from those of T. modestus, syllabized 
a choreéwee repeated over and over, then varied with phrases like cho- 
weéoo and choreéchee, wheeooreé tickwheéoo-wichew (the tick soft); 
also chwee-chweeéo and variations like chwee-chweéeee. Its calls in- 
clude a rather soft trrrup varied to a sharper whrrrp, and a churr; none 
of its calls resemble the very loud rattle of T. modestus. When not 
alarmed they may move actively through low growth, peering about, 
like titmice swinging over to examine the lower surface of limbs and 
twigs. 

Arbib and Loetscher in 1934 (Auk, 1935, p. 327) recorded these 
wrens as “breeding during July and August” in or near the Canal Zone, 
but give no other statement as to their nesting. Skutch (Condor, 1968, 
pp. 70-71) describes nesting of an allied subspecies in Venezuela. He 
found a dormitory as well as a breeding nest, the latter roughly globular 
with an entrance on the side. The occupant was a nestling of the para- 
sitic Shiny Cowbird. In Venezuela this wren often occurs on dry hill- 
sides far from water. 

Stomachs that I examined have been filled with fragments of insects, 
including remains of caterpillars, and with these, bits of spiders and a 
pseudoscorpion. Males weighed by Burton (Bull. Brit. Orn. Club, 
1975, p. 85) range from 18.0 to 23.0 g, females from 16.0 to 19.5 g. 

Lawrence named this race for John R. Galbraith, “an intelligent and 
skillful young taxidermist” from New York, who collected with Mc- 
Leannan along the line of the Railroad in Panama during the winter 
of 1860-61, contributing various notes on habits and other items of 
interest. 


FAMILY TROGLODYTIDAE 7 ip! 


THRYOTHORUS LEUCOTIS CONDITUS (Bangs) 


Thryophilus galbraithi conditus Bangs, Proc. New England Zool. Club, vol. 4, 
March 16, 1903, p. 3. (“San Miguel Island, Gulf of Panama” = Isla del Rey, 
Archipiélago de Las Perlas, Panama.) 


Characters.—Similar in general to T./. galbraithu but darker brown 
on the dorsal surface, sides, lower abdomen, and undertail coverts; 
slightly larger. 

Measurements.—Males (10 from Islas del Rey and Cafias), wing 
64.1-70.8 (67.0), tail 41.7-46.0 (44.0), culmen from base 19.1-21.2 
(20.3), tarsus 23.4-25.3 (24.5) mm. 

Females (10 from Islas del Rey and Cafias), wing 60.2-66.5 (63.3), 
tail 39.0-44.2 (41.6), culmen from base 18.2-20.8 (20.1), tarsus 23.4- 
25.0 (24.2) mm. 

Resident. Found on Islas Rey, Viveros, Puercos, and Cafias, Archi- 
pielago de las Perlas. 

This subspecies, marked by its definitely darker coloration, especially 
on the back, was described from 8 specimens collected by W. W. 
Brown, Jr., near the town of San Miguel, on Isla del Rey in April and 
May, 1900. On first examination of the collection, Bangs (Auk, vol. 
XVIII, January, 1901, p. 30) listed them as “Thryophilus galbraithi,” 
noting only that they were “almost imperceptibly more reddish on the 
back than mainland birds.’ Later, in more careful comparison, he 
formally named the race. The distinction was verified through a second 
series of 10 specimens collected by Brown on another visit in February 
and March, 1904 (Thayer, J. E., and Bangs, O., Bull. Mus. Comp. 
Zool., vol. 46, 1905, p. 154). H. Rendahl (Ark. f. Zool., vol. 13, no. 4, 
1920, p. 43), in a report on a collection of birds made in the Pearl Is- 
lands in 1882 by Carl Bovallius, listed 4 males taken (April 8 to 11) 
on Isla Viveros, adjacent to Rey on the northwest. 

From January 21 to 23, 1960, I found these wrens common on Isla 
Canas, adjacent to Isla del Rey on the east, and collected a series of 3 
males and 4 females. 

As our launch lay at anchor in a sheltered bay on the south side of 
the island on the evening of January 21, I heard an occasional song 
that I attributed to this bird. And the following morning we found 
them common through the rather low open forest, as well as in denser 
growth adjacent to a mangrove swamp. The clear, ringing song, heard 
constantly, was a rapid repetition of a single phrase. The birds were 
common also, but more wary, on the eastern side of Isla del Rey, at 


72 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


Ensenada below Punta de Pena Tiburon, January 24; at Manzanillo, 
above Punta Gorda, farther south, June 25; and along the Rio Cacique 
at the west side of the large bay on the southern side of Rey on Janu- 
anyc7. 

Eisenmann heard a wren, evidently of this genus, singing near a 
stream on Saboga Island in the Perlas on July 9, 1964. He did not see 
the bird. Though unreported from this island, no other Thryothorus 
is known from this archipelago. 


THRYOTHORUS MODESTUS Cabanis; Plain Wren, Cucarachero Sencillo 


Thryothorus modestus Cabanis, Journ. f. Orn., vol. 8, “Nov. 1860” (published 
May 30, 1861), p. 409. (San José, Costa Rica.) 


Rather small; upper surface from crown to back of tail grayish 
brown. Similar in general to Thryothorus leucotis, but with dark bar- 
rings on wings much less prominent. In nominate modestus and elutus, 
distinctly buff on sides, undertail coverts, rump, and wings; one very 
grayish race (zeledont). 

Description.—Length 125-135 mm (elutus), 145-150 mm (zeledoni). 
Adult (sexes alike), crown and hindneck dark, slightly brownish gray, 
becoming olive-brown on back, scapulars, wing, and rump; upper tail 
coverts faintly paler; tail russet or grayish brown, barred heavily with 
dull black; secondaries and outer webs of inner primaries dull rufous 
or grayish brown, barred with black to dusky; primaries otherwise dull 
black; a broad white line above eye; lores and space around eye white, 
with a dull black spot in front of eye, extending in a narrow line be- 
hind; throat white; breast and upper abdomen white or shaded gray, 
tinged with cinnamon-buff on sides; flanks and undertail coverts cin- 
namon or buffy olive; underwing coverts white. 

Immature, somewhat duller, with markings on side of head less 
distinct. 

The Plain Wren occurs from Chiapas, Mexico, through Central 
America to central Panama. The nominate race, modestus, is found 
from central Mexico south through much of Costa Rica, replaced in 
Panama by the subspecies elutus (which ranges north into the extreme 
southeastern end of Costa Rica). Zeledon’s Wren, zeledoni, described’ 
beyond, is a Caribbean form with a range on the Caribbean slope from 
northern Bocas del Toro north in eastern Costa Rica to southeastern 
Nicaragua; currently regarded as a geographic race of T. modestus, it 
is included (with reservations) under that species. 


FAMILY TROGLODYTIDAE 73 


THRYOTHORUS MODESTUS ZELEDONI (Ridgway ) 


Thryophilus zeledoni (Lawrence Ms.) Ridgway, Proc. U.S. Nat. Mus., vol. 1, 
1878, p. 252. (Pacuare, Limon, Costa Rica.) 


Characters.—Similar to Thryothorus modestus elutus (Bangs), but 
larger and decidedly darker; dorsal surface, sides, flanks, and undertail 
coverts gray (without definite reddish brown markings); bill and feet 
larger, stronger. 

Kennard recorded a female taken at Almirante as having the maxilla 
black; mandible yellow; iris gray-brown; tarsus dark olive. 

Measurements.—Males (10 from Bocas del Toro, Costa Rica, and 
Nicaragua), wing 62.3-65.0 (63.5), tail 49.4-53.9 (51.5), culmen from 
base 19.5-22.5 (21.1), tarsus 24.8-26.6 (25.8) mm. 

Females (9 from Bocas del Toro and Costa Rica), wing 60.0-63.3 
(61.7), tail 48.5-51.5 (48.6), culmen from base 17.4-21.5 (19.9), tar- 
sus 24.2-25.8 (25.0, average of 8) mm. 

Resident. Recorded in the lowlands of western Bocas del Toro, 
from Almirante, Isla Colon in Almirante Bay, and Zegla on the Rio 
Changuinola to the Costa Rican boundary on the lower Rio Sixaola; 
not abundant. 

This wren was described originally by Ridgway as a distinct species, 
from its larger size and darker coloration compared to Thryothorus 
modestus, and was so treated by Salvin and Godman in 1880, Sharpe in 
1881, Carriker in 1910, Kennard and Peters in 1931. Hellmayr, how- 
even, (Cat. Birds Amer., pt. VI, 1937, pp. 170-171), from examina- 
tion of 4 specimens from Matina, Limon, and La Iberia Farm on the 
base of Volcan de Turrialba, Costa Rica, placed it as a race of Thryo- 
thorus modestus with the statement that though “well characterized by 
much larger feet and bill, much duller and less brownish upper parts 
with brownish instead of rufescent wings and tail, and much less ful- 
vous flanks and undertail coverts . . . is clearly conspecific with T. 
modestus . . . though intergradation, especially in dimensions, is far 
from being complete.’’ Hellmayr’s action has been followed, mainly 
uncritically, since. 

Carriker (Ann. Carnegie Mus., vol. 6, no. 4, 1910, p. 755), in writing 
of Costa Rica, said of geledoni that he first encountered it “along the 
Sicsola [=Sixaola] River, where they frequented wild-cane brakes and 
a high tangled grass found only along the river bank, known as ‘gami- 
lote.’” He never saw it in forest. His locality was along the present 
northeastern end of the boundary with Panama. 


74 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Karly reports in Panama were by Griscom (Amer. Mus., Novy. no. 
293, 1928, p. 1) of 1 collected near Almirante by Benson, and by Ken- 
nard and Peters (Proc. Boston Soc. Nat. Hist., vol. 38, no. 10, 1928, p. 
459) of 1, taken also at Almirante, by Kennard, on March 16, 1926. 
Peters later (Bull. Mus. Comp. Zool., vol. 71, 1931, p. 330) recorded 
a female from this locality collected by Hasso von Wedel, August 10, 
1927. A male and female in the Havemeyer Collection at the Peabody 
Museum were taken there by Austin Smith, April 19 and 23, 1927. 

Eisenmann (Condor, 1957, p. 256), who observed these birds at 
Changuinola in 1956, with reference to Hellmayr, described the song 
as loud and ringing and wrote, “. .. those that I heard from zeledom 
were different from any I have noted from elutus, but they showed 
some resemblance.” Slud (Amer. Mus. Nat. Hist., Bull. 128, 1964, p. 
285) who described the notes in detail, concluded that “the songs of 
modestus and zeledoni sound identical to me except that zeledoni gives 
....an uncharacteristic deliberate swir, swirrst, the last note rising, so 
to speak, in an overhanging curve. Also zeledoni has a harder, heavier 
call note.” He considered that birds from the Terraba Valley, on the 
Pacific slope in the southwest, showed approach to zgeledom in bill size 
and grayness. Dr. Thomas Howell, who has seen the bird in Nicaragua, 
informs me (i litt.) that to him the song and calls suggested those of 
T. modestus. 

My own personal knowledge of zeledomi has come from encounters 
near Changuinola in western Bocas del Toro, where I collected 1 
February 13, 1958, and saw others briefly February 28, but did not 
hear the calls or song. Museum specimens have increased, so that from 
the 4 listed by Hellmayr in his discussion on relationship, | have mea- 
surements of 7 males and 5 females (including Ridgway’s type), and 
have examined several others in which the sex has not been marked. All 
have shown the larger size and the darker, distinctly gray coloration on 
which Ridgway described the bird as a distinct species, with no inte- 
gration to the smaller size and brighter colors of T. modestus elutus 
with which it has been supposed to hybridize. From my examination 
of specimens it has seemed to me that geledoni is a distinct species. 
However, because of my slight personal knowledge of the bird in life it 
is listed here as a geographic race, with the suggestion that it be studied 
further when there is suitable opportunity. 

The range of geledomi, from the published record in Panama, is 
restricted to the Caribbean lowlands, inland only to the interior foot- 
hills. In Costa Rica, it ranges only on the Caribbean coastal plain 
(being replaced by nominate modestus at about 600 m, fide Slud, 


FAMILY TROGLODYTIDAE 75 


1964, op. cit.), continuing to southeastern Nicaragua, at Los Sabalos 
where the river of that name joins the Rio San Juan, at San Juan del 
Norte (formerly Greytown) at the mouth of this river. 

The eggs of T. m. geledoni have not as yet been described. A nest 
collected at Almirante June 4, 1962, by Pedro Galindo is an elliptical 
ball of long slender plant fibers (apparently aerial rootlets, mixed with 
fine grass stems and a lesser amount of broad-bladed grass) measuring 
120X160 mm. The entrance hole in the end was underneath so that it 
was sheltered. Another nest, slightly smaller, with dimensions 110 
120 mm, found on the same date, was more loosely compacted of 
shorter, coarser materials, including rootlets and grass. It also had the 
entrance hole sheltered beneath, at one end. Both nests were empty. 


THRYOTHORUS MODESTUS ELUTUS (Bangs) 


Thryophilus modestus elutus Bangs, Proc. New England Zool. Club, vol. 3, Janu- 
ary 30, 1902, p. 51. (Lion Hill Station, Panama Railroad, Canal Zone.) 


Characters.—Smaller; rump and upper tail coverts russet, tail dull 
russet; upper abdomen and tibia white, changing to gray on upper sides; 
lower sides, flanks, and undertail coverts buffy cinnamon. 

A male at El Copé, Coclé, February 24, 1962, had the iris reddish 
brown; maxilla black; nasal operculum mandible, tarsus, toes, and 
claws dark neutral gray. Posterior area of the inside of the mouth, in- 
cluding the thickened base of the tongue, black, shading anteriorly to 
neutral gray on inner surface of bill. The slender tip of the tongue was 
transparent, without color. In another male, at Puerto Armuelles, 
Chiriqui, March 10, 1966, the iris was orange-brown; tip of mandible 
dark neutral gray; cutting edge of maxilla and base of mandible light 
neutral gray; rest of maxilla black; tarsus, toes, and claws dusky neutral 
gray. 

Measurements.—Males (10 from Chiriqui, Veraguas, Los Santos, 
Herrera, Coclé, and Canal Zone), wing 58.1-60.9 (59.2), tail 45.6- 
51.2 (48.2), culmen from base 18.1-18.9 (18.1), tarsus 23.0-24.8 
1235-0) mim. 

Females (10 from Chiriqui, Veraguas, Los Santos, Coclé, and Canal 
Zone), wing 53.4-56.7 (55.2), tail 42.0-49.5 (44.2), culmen from base 
17.1-18.9 (18.1), tarsus 21.6-23.0 (22.6) mm. 

Resident. Common in the Tropical Zone of the Pacific slope; from 
western Chiriqui (to over 1500 m above El Volcan including Cerro 
Punta; above Boquete) and the Burica Peninsula at Puerto Armuelles, 
east through southern Veraguas, the Azuero Peninsula to Punta Mala, 


76 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


the western and eastern sections of the Province of Panama, the lower 
Bayano Valley (Chepo, San Antonio); in the northern Canal Zone 
also on the Caribbean slope, including the lower and middle Chagres 
Valley (Juan Mina, Lion Hill, Gatun, Fort San Lorenzo) and the Rio 
Boqueron above Madden Lake (Peluca Hydrographic Station) and the 
Province of Colon east to Portobelo. It follows clearings into the 
mountains (reaching 1860 m at Cerro Punta) of Chiriqui (Rio Chi- 
riqui, Fortuna Dam site), Veraguas (Santa Fé, Chitra), Panama 
(Cerro Campana, Cerro Azul-Cerro Jefe area). 

From the lowlands of western Chiriqui the subspecies elutus 
ranges for a short distance into southern Costa Rica through the Golfo 
Dulce area to the lower valley of the Rio Terraba (specimens in collec- 
tions of the Western Foundation of Vertebrate Zoology). When found 
in the same general area as the more secretive Buff-breasted Wren 
(Thryothorus leucotis galbraithu) the Plain Wren may be identified by 
its paler colors and less heavily barred wings. 

These wrens are found in thickets, brushy pastures, and low cover 
in other open areas, including suburban gardens, hedgerows, and weed- 
grown fields, outside the heavy forests. Here they forage in pairs, and — 
at the proper season, in family groups. While usually encountered low 
down, they also explore the branches of trees above the thickets, espe- 
cially when these have tangles of vines that provide cover. In such lo- 
cations they move about with much low chatter, often concealed among 
the leaves, hiding carefully at any alarm. Neglected orange and grape- 
fruit groves, where the trees have many epiphytes, are often attractive 
to them. 

In Eisenmann’s opinion, elutus is, after the House Wren, the most 
numerous wren in open, extensively deforested areas of Panama, chiefly 
in the Pacific lowlands, but also very common in the Boquete district, 
Chiriqui, and the cleared areas of the Caribbean watershed in the Canal 
Zone. The highest bird he has found was seen singing in a field at 
Cerro Punta, Chiriqui, at 1860 m on December 11, 1962. While un- 
recorded in Darién, where replaced by the essentially South American 
T. leucotis, both species occur sympatrically in the Canal Zone and in 
open or cleared areas of eastern Province of Panama. 

The song, heard constantly, is a series of whistled notes, ending 
with one or two in harsher tone, from which country residents in Costa 
Rica call them “chinchirigui” (varied by some to “richichi”’). When 
heard near at hand this is found to be a combination of three or more 
whistled sounds followed by a harsher call, the first given by the male, 


FAMILY TROGLODYTIDAE I 


the second immediately following by the female. They also have the 
rattling notes common to related wrens. Skutch (Pacific Coast Avif., 
no. 34, 1960, p. 125) describes a varied series of musical sounds that he 
attributed to immature birds. 

Eisenmann considers the songs, although musical and varied, to lack 
the richness and sweetness, and often the loudness of those given in 
central Panama by other wrens of the genus Thryothorus. Most songs 
include the cheénchirigweé phrase that gives the species its vernacular 
name in Costa Rica. Sometimes the last note slides down at the end, 
or the phrase may be changed to weereeweé; these phrases are often 
introduced by other notes, or the phrase may be replaced by a more 
musical cheechooee or cheeweetweet or variants. A very distinctive call 
is a loud, rapid tsiptsip-tsrrrrrr, sometimes just tsirrrrrr, suggesting 
the sound produced by the whirling wooden rattle often used in New 
Year’s Eve celebrations. At El Valle, Coclé, on August 30, 1951, he 
observed a pair duetting: one delivered a phrase of usually four notes 
and the other added two, over and over (see also Worth, Bird-Lore, 
1939, p. 282 on antiphonal singing in Chiriqui by birds perched a few 
inches from each other). 

Blake (Condor, vol. 58, 1956, p. 387) recorded a nest and eggs, col- 
fecred by P2 Monniche near. Boquete, May 10, 11932, as the usual 
rounded ball “loosely woven of dried grass” with a lining of finer ma- 
terial and chicken feathers. The two partly incubated eggs were “dull 
white, unmarked, and measure 21.5 14.3 and 22.314.2 mm.” The 
field journal of G. Ralph Meyer for July 7, 1941, records a nest in the 
Canal Zone built in a croton hedge along the front of the military quar- 
ters. This contained three white eggs which seemed very large for the 
size of the bird. As they were well incubated they were not collected. 

Six nests in the collection of the Western Foundation of Vertebrate 
Zoology, collected by Andrews Williams and Mrs. Elsie Fiala from 
March 21 to June 30, 1971, in the Rio Terraba-Golfo Dulce area in 
Costa Rica, were alike in rounded form, with the entrance at one side. 
Each held two eggs. One, on April 10, 1971 (parent wren flushed from 
the nest), with the entrance on the side near the top, held one egg of the 
wren and one of the parasitic Striped Cuckoo, Tapera naevia excellens. 

Skutch (Pac. Coast Avif., no. 34, 1960, pp. 123-129) describes be- 
havior and nesting, chiefly in Costa Rica, pointing out that in addition 
to the globular breeding nest, this species builds a domed dormitory 
nest of much looser construction with a larger side entrance, used by the 
male while the female sleeps in the breeding nest. 


78 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


THRYOTHORUS RUFALBUS CASTANONOTUS (Ridgway): 
Rufous-and-white Wren, Cucarachero Rojizo 


Thryophilus rufalbus castanonotus Ridgway, Proc. Boston Soc. Nat. Hist., vol. 
XXIII, March, 1888, p. 386. (“Nicaragua to the highlands of Columbia” = 
Angostura, Cartago, Venezuela.) 


Medium size: above chestnut-brown, a prominent white line above 
the eye; undersurface white, except for brownish flanks. 

Description Length 135-155 mm. Adult (sexes alike), upper sur- 
face pale to deep chestnut (lighter on crown); wings narrowly, tail 
more broadly, barred with black; a distinct white superciliary line, 
margined narrowly above by black; line on side of head from the eye 
to neck rufous-brown; rest of side of head white, streaked with black, 
with lower side of head white, bordered below by a narrow black line; 
undersurface white, changing to grayish brown on sides, browner, be- 
coming darker on flanks; underwing coverts spotted with black; under- 
tail coverts white, barred heavily with black, the dark bars edged nar- 
rowly with rusty brown. 

Immature, duller, darker brown above, especially on crown and hind- 
neck; duller white on lower surface, with breast lined distinctly with 
dusky; undertail coverts pale rusty brown, barred lightly with black. 

An adult male, collected at Chiva Chiva, Canal Zone, March 19, 1961, 
had the iris dark brown; cutting edge of maxilla and mandible dull 
whitish; rest of maxilla fuscous-black; base of gonys and anterior half 
of mandible (except cutting edge) pale neutral gray; tarsus, toes, and 
claws brownish gray. Another male at Las Palmitas, Los Santos, Janu- 
ary 24, 1962, had the iris dark reddish brown; maxilla and tip of mandi- 
ble black; rest of mandible light neutral gray; tarsus, toes, and claws 
light mouse brown. 

Measurements.—Males (10 from Chiriqui, Veraguas, Los Santos, 
Herrera, Coclé, Province of Panama, and Canal Zone), wing 67.2- 
72-1 (69. 3), tail 48.2-54.7 (52.1), culmen from base 19.7-23.6 (21.3), 
tarsus 24.2-25.7 (25.0) mm. 

Females (10 from the same localities as the males), wing 61.2-67.8 
(64.3), tail 46.2-51.5 (48.4), culmen from base 19.5-21.5 (19.7), tarsus 
23.8-25.0 (24.5) mm. 

Resident. Widely distributed on the Pacific slope in the lowlands, 
where it is locally common in thickets and deciduous woodland, from 
western Chiriqui to the Rio Bayano Valley, following clearings into the 
highlands, and on the Caribbean side of the Canal Zone (especially mid- 
dle Chagres Basin and adjacent Colon Province, but rather rare near 


FAMILY TROGLODYTIDAE 79 


the Caribbean coast. In Chiriqui, from Quebrada Santo Domingo 
(west of Concepcion and Alanje) east through David to San [elix; 
Veraguas, including Remedios, Cerro Flores, Chitra, Santa Ié, Calove- 
vora, and Santiago as well as the western side of the Azuero Peninsula 
(Paracoté, Mariato River); Los Santos (to Punta Mala); Herrera; 
southern Coclé; western Province of Panama (Playa Coronado, Cerro 
Campana), and the eastern section to La Jagua, Chepo, San Antonio, 
and the lower Bayano Valley. In the Canal Zone found mainly on the 
Pacific side near Panama City and Chiva Chiva; Goldman, in 1912 
secured 1 at Tavernilla, now submerged in Gattn Lake near Barro 
Colorado Island. Major General Meyer found it near Summit. In 
mountain areas it is recorded to 750 m near El Valle, Coclé, and to 1000 
m on Cerro Hoya, Los Santos, and to 1500 m at Boquete (Quiel). 

These birds are found in pairs, usually on or near the ground in 
thickets of fairly open areas. Often they appear less timid than most 
other species as they range in brush-filled gullies or low thickets in 
open areas, or in the denser ground cover in gallery forest. They seem 
adaptable, and are able to exist in second growth when this follows 
clearing of the original forest. As they move about under cover of vines 
and low shrubbery, their presence may be known from the songs of the 
males. These begin with a rolling repetition of a loud note that changes 
to a trill, the two mingling with varying cadence in pleasing combina- 
tion. On the whole, they sound less emphatic in utterance than the 
songs of others of the genus found in Panama. Songs attributed to the 
females, heard alternately with those of males, are softer, less emphatic 
in tone. They often respond to whistled imitations, and also are at- 
tracted by squeaking sounds. 

The field notes of Major General G. Ralph Meyer for July 30, 1941, 
describe a nest, found near the Gamboa Road junction, as bulky, made 
of grass, hanging in the fork of a “bull thorn” or bullhorn acacia (Aca- 
cia costaricensis and allies). The tubular entrance was almost at a right 
angle to the axis of the main nest, which lay along the trunk of the tree. 
The whole was about 300 mm long by 125 mm wide, with the tubular 
entrance about 50 mm in diameter. Other nests were present in the 
same kind of small tree, infested with fierce small ants (Pseudomyrma) 
that live symbiotically on these plants. Meyer recorded that this wren 
was host to the parasitic Striped Cuckoo, Tapera naevia excellens. (See 
Wetmore, Smithsonian Misc. Coll., vol. 150, Part 2, 1968, p. 135). 
Meyer described the three eggs found in the nest as colored greenish 
white, with measurements as follows: 21.6 15.7, 22.2 15.2, and 22.3 x 
bo./ mm. 


80 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


Two males collected by Strauch (Bull. Brit. Orn. Club, 1977, p. 64) 
weighed 27.1 and 27.7 g. 

Although primarily a lowland species, Eisenmann (in litt.) saw it at 
about 1500 m in the Boquete area at Quiel in a coffee plantation (Febru- 
ary 27, 1960) and on the Camino a la Estrellon (July 19, 1964, seen 
singing). Ridgely netted and released 1 on March 3, 1976, at approxi- 
mately 1350 m in central Chiriqui on the upper Rio Chiriqui (Fortuna 
Dam site) at the edge of a clearing in humid forest, a surprising 
location. 

The song, while variable, is usually very distinctive and unlike that of 
any other wrens because of the low pitch of many notes, which, to 
Eisenmann’s ear, gives a “hooting” quality, although Hartshorne states 
they are pure, musical tones, mostly on the diatonic scale. The most 
characteristic phrase consists usually of four (more or less) “hooting” 
notes often followed by an emphatic higher note: hoo-hoo-hoo whit; 
sometimes the higher last note slides down or is followed, or replaced 
by, a trill. Sometimes the initial notes (lee, lee, looo) suggest the first 
three notes of the bugle call “taps.” Mrs. Cora C. Alderton, who lived 
at Gamboa, Canal Zone, had a nesting pair in her garden in 1957-58 
and gave Eisenmann, in musical notation, fifteen major variations and 
a larger number of minor ones of songs heard, and said she had heard 
more than fifty variations. Among, the “trilling”’ variations is one 
somewhat resembling a subdued bugle call, mentioned by Sturgis (Field 
Book of Birds of the Canal Zone, 1928, p. 358). The song has been 
highly praised also by Chapman (My Tropical Air Castle, 1929, p. 358) 
and others. In addition to the songs, Eisenmann has heard a loud, 
rattling chatter and a dry thrrrrp. 

Mrs. Alderton showed Eisenmann a nest in her garden in Gamboa 
that was “saddle bag-shaped,” with a wide entrance tube, on the stem 
of a frond of a small ornamental palm. It was begun on April 3, 1958, 
but destroyed by Thraupis tanagers. Another nest was begun in the 
same palm on April 11; straws for the lining were brought in on April 
27. Incubation apparently began in early May. D. E. Harrower (thesis 
at Cornell University) reports “breeding season from at least April to 
September’; he found an active nest on July 17 at Pedro Miguel, Canal 
Zone, in a thorny acacia 2-3 m from the ground. F. O. Chapelle noted 
a pair building near Ft. Clayton on April 20, 1955; this nest, also in 
bullhorn acacia, was almost finished on April 23. S. F. Ambrose found 
a nest being built in a bullhorn acacia on July 4, 1964, at Santa Clara, 
Coclé, on the Pacific Coast. 


FAMILY TROGLODYTIDAE SI 


A bird banded by H. Loftin at Curundu, Canal Zone, on December 
7, 1963, was recaptured there on March 5 and April 30, 1966, and on 
September 20, 1967. 


THRYOTHORUS THORACICUS Salvin: Stripe-breasted Wren, 
Cucarachero Pechirayado 


Thryothorus thoracicus Salvin, Proc. Zool. Soc. London, 1864, p. 580. (Tucurrique, 
Costa Rica.) 


Rather small; dull brown above; foreneck and breast white, heavily 
streaked with black. 

Description.—Length 106-122 mm. Adult (sexes alike), above 
brown, varying from a dull grayish brown hue on forecrown to russet- 
brown on rump and upper tail coverts; in some individuals, very faintly 
and indistinctly barred narrowly with black; wings and tail somewhat 
lighter brown, heavily barred with black; side of head and of upper 
neck dull brownish black streaked with white; a white superciliary line; 
foreneck and breast white, lined broadly with black, these markings re- 
duced in extent on abdomen; sides grayish olive, changing to rusty 
brown on flanks; undertail coverts brown, barred with black; under- 
wing coverts white, barred and lined with grayish brown. 

Immature, undersurface lighter, somewhat duller brown with black 
lines greatly reduced in extent, less definite. 

Measurements.—Males (10 from Veraguas and Costa Rica), wing 
55.0-62.5 (59.0), tail 37.1-40.4 (38.1), culmen from base 18.1-20.0 
(19.0, average of 9), tarsus 20.6-22.3 (21.6) mm. 

Females (10 from Veraguas, Bocas del Toro, Costa Rica, and Nica- 
ragua), wing 55.8-62.1 (58.1), tail 34.4-39.8 (36.8), culmen from base 
17.3-18.9 (18.2), tarsus 19.3-22.0 (20.7) mm. 

Resident. Rare; known on the basis of specimens on the Caribbean 
slope of Panama, from western Bocas del Toro (recorded from Zegla, 
Rio Changuena ), to the border with Veraguas (Rio Calovévora, Amer- 
ican Museum of Natural History) and from Cascajal, northern Cocle. 
There are sight records from the Canal Zone (Gattin Dam, Achiote 
Road) (Ridgely, 1976, p. 267). 

The first authentic published record for the Republic is a male labeled 
“Almirante,” forwarded by Hasso von Wedel, dated May 16, 1927. 
(Peters, Bull. Mus. Comp. Zool., vol. 71, 1931, p. 330). It is of interest 
to note that Austin Smith, in the final days of a trip to Panama in 1927, 
worked briefly in Bocas del Toro. Among his specimens, preserved in 
the Havemeyer Collection in the Peabody Museum in New Haven, are 


82 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


2 males of this wren from Zegla, Rio Teribe, collected May 16, 1927. 
The locality was a small village located near the mouth of the Rio 
Teribe, where that stream enters the Rio Changuinola, about 4 km above 
the bridge on the railroad, north of Almirante. It appears that Wedel, 
cited above, either accompanied Smith or acquired one of his specimens. 

Our collections also include 2 skins of T. thoracicus, collected Sep- 
tember 8 and 14, 1961, by Pedro Galindo at a camp at 750 m elevation 
on the Rio Changuena, a tributary of the Rio Changuinola farther in- 
land. There is no other report from the well-known Almirante area. 

Another record for the Caribbean slope is an adult dated March 3, 
1889, marked Cascajal, Coclé, collected by the missionaries H. F. Heyde 
and Ernesto Lux. The locality is on the Rio Cascajal on the Caribbean 
slope in northwestern Cocle. 

In the American Museum of Natural History are 9 ee, labeled 
from Guaval, Rio Calovévora, Veraguas (266-500 m), taken by R. R. 
Benson between August 1-September 19, 1926. This river forms the 
boundary between Bocas del Toro and Veraguas. One specimen from 
September 4 is a full-grown bird in juvenal plumage. 

E. O. Willis (Eisenmann, im litt.) reports seeing a pair in September 
1963 in the northwestern Canal Zone near the Caribbean coast on the 
road to Rio Medio, following army ants. He writes that the vocalization 
consisted mainly of single, similar notes somewhat resembling a Night- 
ingale Wren but repeated more rapidly and sometimes varied with notes 
of other pitches of a more Thryothorus quality. Pujals also reports 
having once seen this species along the Achiote Road of the northwest- 
ern Canal Zone, and later on February 14, 1947, near Rio Indio village 
in western Colon. 

A record by Salvin (Proc. Zool. Soc. London, 18673;pyia- anes 
ports 3 specimens received from Arcé, erroneously as from “San- 
tiago de Veragua,” far down in the Pacific lowlands of Veraguas. 
This error was repeated by Salvin and Godman (in Biol. Centr.-Amer., 
Aves, vol. 1, 1880, p. 86). The bird is known in Panama almost ex- 
clusively from the Caribbean slope, and so may have been taken in the 
Calovévora area on the northern Caribbean slope of the mountains, ac- 
cessible by a well-known trail from Arcé’s permanent location at Santa 
Fé. On March 18, 1974, Skutch and Eisenmann (in. litt.) heard the 
distinctive call of this species in the humid forest on Santa Fé, on the 
Pacific slope. Evidently the bird does « cross the Divide in at least this 
location. 

Arcé, in the early days of his work in Panama, seems not to have 
labeled some of his specimens carefully. Salvin, with the lack of detail 


FAMILY TROGLODYTIDAE 83 


in geographic data available, then seems also not to have understood 
clearly his collector’s location. In some instances he seems to have in- 
terpreted the shipping point, in the present case Santiago, for the col- 
lector’s location, far distant in the mountains. 

This wren has been studied in detail by Skutch on the humid tropical 
Caribbean slope of Costa Rica (Publ. Nuttall Orn. Club, no. 10, 1972, 
pp. 159-163). The song consists of two types, the first being a distinc- 
tive series of slowly repeated whistled notes, almost like those of a 
pygmy-owl, heard usually at dawn and the second is a medley of sounds 
more usual in the family, highly varied and often antiphonal in utter- 
ance by paired male and female. The ball-shaped nest has two sections, 
an outer part with a broad opening downward, with a passage to the in- 
ner chamber that holds the eggs. These, laid in April—in one case three 
in number, in another, two—are plain white without markings. The 
young at hatching are without down. 

Carriker (Ann. Carnegie Mus., vol. VI, 1910, p. 759) found a nest 
at Jimenez, Costa Rica, May 9, 1905, with three immaculate white eggs, 
so heavily incubated that only one was preserved. This measured 19.5 
x14 mm. 

My only personal acquaintance with this species came in November 
1940 on the Caribbean slope of the Continental Divide on Cerro Santa 
Maria in northern Costa Rica. Here its clear songs were heard regu- 
larly, usually about deadfalls in heavy forest. 


THRYOTHORUS LEUCOPOGON GRISESCENS (Griscom): 
Stripe-throated Wren, Cucarachero Garganta Rayada 


Thryophilus leucopogon grisescens Griscom, Bull. Mus. Comp. Zool., vol. 72, 
January 1932, p. 359. (Permé, San Blas, Panama.) 


Rather small; dull brown; side of head black, streaked with white; 
throat dull white, streaked indistinctly with dusky; rest of undersurface 
plain, without markings, undertail coverts more rufescent. 

Description.—Length 115-125 mm. Upper surface, including scapu- 
lars, dark grayish brown, very faintly and indistinctly banded with 
black; wings and tail light chestnut-brown banded narrowly with black; 
middle and lesser wing coverts slightly paler than back, banded nar- 
rowly with dull black; lores centrally dull black, above white, this color 
extending back as a line over eye and auricular area, bordered above by 
a narrow black line that, as a small black spot, breaks the white at the 
center of the upper eyelid; side of head black lined with white with a 
black line along lower jaw; throat dull white, with dusky; undersurface 
dull tawny brown, slightly paler in the center of the abdomen, more 


84 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


rufescent on the undertail coverts and duller on the sides; axillars and 
underwing coverts dull white, mixed indistinctly with dusky. Young 
birds have the throat buffy with the dark lines very indistinct. 

A male taken at Armila, San Blas, February 26, 1963, had the iris 
auburn, narrow cutting edge of maxilla dull white; rest of maxilla 
black; mandible dark neutral gray on sides, neutral gray on lower sur- 
face; tarsus and toes fuscous-brown; claws dark neutral gray. An- 
other male at this locality, March 2, 1963, had the iris orange; tarsus 
and toes brownish neutral gray. 

Measurements.—Males (10 from San Blas and Darién) wing 55.4- 
58.0 (56.7), tail 34.3-39.7 (37.1), culmen from base 17.5-19.3 (18.5), 
tarsus 18.0-21.7 (20.4) mm. 

Females (6 from eastern Province of Panama, San Blas, and Da- 
rién), wing 51.6-56.2 (53.1), tail 34.4-36.2 (35.3, average of 4), culmen 
from base 17.3-19.3 (17.9, average of 5), tarsus 18.6-20.2 (19.3) mm. 

Resident. Found locally in eastern Panama; on the Caribbean slope 
in eastern San Blas from Permé and Armila to the Colombian boundary 
at Puerto Obaldia; on the Pacific side, from Cerro Chucanti, at the head 
of Rio Majé, in extreme eastern Province of Panama, and Darién (re- 
corded at Tapalisa, Cituro, Pucro, and at the base of Cerro Quia (Rio 
Mono) in the upper Tuira Valley, and near Cana, on Cerro Pirre; near 
Jaqué and on the upper Rio Jaqué). Outside of Panama, this species is 
found in western Colombia and northwestern Eucador. 

These wrens range more in the open than most of their relatives. 
While they may be found low down, in dense growth near the ground, 
they also are regularly seen higher, among vines and branches in forest 
areas. The only one seen on Cerro Chucanti associated with a group of 
small ant-wrens in the lower branches of open forest. One near Armila 
hunted through the higher limbs in a cacao plantation, and another on 
the Rio Pucro was in open cover near a trail leading through a forest. 
On the Rio Jaqué, 1 searched through a mass of vines 10 m above the 
ground, and at Jaqué I found 1 in the undergrowth in heavy forest. At 
other localities I saw them regularly in fairly open areas at heights of 
15 to 18 m. 

At Armila, on March 2, 1963, a pair was busily building a nest in 
fairly open forest at the border of a quebrada. The structure was an 
untidy ball, irregular in form, placed near the end of small branches 
with no attempt at concealment. 

Hellmayr (Cat. Birds Amer., pt. 7, 1934, p. 176), with a rather small 
series, listed the Thryothorus leucopogon group as a geographic race 
of the widely separated T. thoracicus, a treatment that has been ac- 


FAMILY TROGLODYTIDAE 85 


cepted by other recent students. With the considerable series now avail- 
able it is clearly evident that thoracicus is uniformly and _ heavily 
streaked with black and white over throat and breast. In T. leucopogon 
the marking is confined wholly to the upper throat, with foreneck and 
breast immaculate. The two styles of marking are clear cut in their 
differences, with no intergradation. 


THRYOTHORUS NIGRICAPILLUS Sclater: Bay Wren, 
Cucarachero Cabecinegro 


Ficure 10 


Thryothorus nigricapillus P.L. Sclater, Proc. Zool. Soc. London, pt. 28, May 
1860, p. 84. (Nanegal, 4000 feet elevation, Ecuador.) 


Medium size; crown and hindneck black, rest of upper surface 
chestnut-brown, with wings and tail barred black; throat white, rest of 
undersurface chestnut or barred. 


Ficure 10—Bay Wren, Cucarachero Cabecinegro, Thryothorus nigricapillus. 


86 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Description.—Length 126-156 mm. Crown and hindneck black in 
sharp contrast to chestnut-brown of back and upper tail coverts; wings 
and tail somewhat lighter brown, heavily barred with black; throat, 
line over the eye, lores, auricular region, and side of jaw white; a nar- 
row black line below along the side of the jaw. The pattern of color on 
the upper surface in all of the races is as described above. The under- 
surface differs widely in the subspecies, from plain chestnut-brown in 
costaricensis and odicus, to white, barred narrowly with black on the 
undersurface, in the eastern race schottw (in which there are two dark » 
bars on each feather). 

The range of this species, as here conceived, extends along the Carib- 
bean slope of Nicaragua, Costa Rica, and Panama (both slopes from 
Veraguas eastward), western Colombia, and northwestern FEucador. 
(See Wetmore, Smiths. Misc. Coll., vol. 139, no. 2, 1959, pp. 14-17 for 
taxonomic review). 

The call note in all is a somewhat guttural chur-r-rk. Song utterances 
are varied, but all agree in their ringing quality and cadence. The 
usual variants that may be heard from one male in the course of a few 
minutes I have written as follows: commonly, chit-i-sit chit-1-sit, re- 
peated rapidly a dozen times, then perhaps varied to sweety-it sweety-it. 
This may change to sweety-tu sweety-tu, or sweet-whee-hee sweet- 
whee-hee. The changes form quite a repertoire, but in final analysis all 
are rather similar. Males sing alone, but antiphonal duetting by a pair 
is more common. 


THRYOTHORUS NIGRICAPILLUS COSTARICENSIS (Sharpe) 


Thryophilus costaricensis Sharpe, Cat. Birds Brit. Mus., vol. 6, 1881, p. 217. 
(Costa Rica = Valley of the Rio San Carlos, Alajuela, Costa Rica.) 


Characters.—Ear coverts, side of jaw, throat, and upper foreneck 
white; rest of undersurface auburn to hazel; some with sides barred 
indistinctly with black. 

Measurements.—Males (10 from Bocas del Toro), wing 66.5-71.5 
(69.6), tail 51.1-56.8 (54.4), culmen from base 20.4-22.4 (21.3), tarsus 
24.6-26.9 (25.7) mm. 

Females (10 from Bocas del Toro), wing 62.5-67.2 (65.2), tail 46.3- 
51.3 (49.2), culmen from base 19.7-21.8 (20.7), tarsus 24.1-27.2 
(25.2) mm. 

Resident. Common in the lowlands of Bocas del Toro, from the Rio 
Sixaola to the seaward side of the base of the Valiente Peninsula 
(Cocoplum). 


FAMILY TROGLODYTIDAE 87 


This common lowland wren, found in thickets in swampy, low-lying 
areas on the shores of Almirante Bay, is known mainly by its songs 
and chattering, and scolding notes, since the birds remain hidden in 
dense cover. They were observed regularly in pairs. 

Carriker (Ann. Carnegie Mus., vol. 6, 1910, p. 761) reports finding 
many nests in the Caribbean lowlands of Costa Rica, including the Rio 
Sicsola [= Sixaola], the boundary with Panama in Bocas del Toro. 
The nest is an elbow-sloped structure usually hung in an upright crotch 
of some small tree or shrub from 1.6 to 5 m above the ground, generally 
about 25 cm long and 8 to 13 cm in diameter at the larger end. All Car- 
riker found were built of weed stems, grass, rootlets, and skeletonized 
leaves, and lined with soft skeletonized leaves. Two nests, each with 
three fresh eggs on May 9 and 11, apparently contained the full com- 
plement. The eggs were “pure white, sparingly speckled over the whole 
surface (more thickly about the large end) with cinnamon-brown.” 
One set measured 23X16, 2516.5, and 24X17 mm. 

The type of this race, named by Sharpe, in the British Museum, is 
listed in the volume on type specimens (Warren and Harrison, vol. 2, 
Passerines, 1971, p. 132), as “Costa Rica. Collected by and purchased 
of A. Boucard.” The collector (Proc. Zool. Soc. London, 1878, p. 51), 
who listed the bird under the name “Thryophilus castaneus, Lawr.,” 
said of it “Several specimens from San Carlos; killed in February.” 
He described the locality (cit. supra, p. 38) as on Rio San Carlos, a 
tributary of Rio San Juan on the Caribbean slope. I have, therefore, 
designated this area as the type locality (Wetmore, Smiths. Misc. Coll., 
vol. 139, no. 2, 1959, p. 19). 


THRYOTHORUS NIGRICAPILLUS ODICUS Wetmore 


Thryothorus nigricapillus odicus Wetmore, Smiths. Misc. Coll., vol. 139, no. 2, 
July 8, 1959, p. 14. (Isla Escudo de Veraguas, off the base of the Valiente 
Peninsula, Bocas del Toro, Panama.) 


Character.—Similar to Thryothorus n. costaricensis, of the adjacent 
mainland, but definitely larger; bill longer, heavier; brown of back and 
undersurface definitely paler, ochraceous-tawny. 

Measurements.—Males (5 specimens), wing 75.2-79.2 (77.0), tail 
98.6-62.3 (60.2), culmen from base 21.8-24.2 (23.2), tarsus 28.4-31.8 
(29.7) mm. 

Females (6 specimens), wing 70.2-72.8 (71.6), tail 54.5-58.8 (56.8), 
culmen from base 21.0-22.3 (21.5), tarsus 26.1-28.7 (27.2) mm. 

Resident. Found only on Isla Escudo de Veraguas, Bocas del Toro. 


88 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


When I visited the island, this wren, most common of the resident 
land birds, lived in pairs scattered through the undergrowth. In this 
island home, with no human neighbors, the birds were tame and easily 
observed. Though most often encountered in low tangles where vines 
were matted and cover was dense, they ranged also into open areas 
through the thickets, occasionally even overhead into the higher 
branches of the forest trees. The song, heard constantly, resembled 
that of the mainland race, but was slightly higher in tone and somewhat 
less varied in repertoire. Regularly the birds came fearlessly near at 
hand, even within a distance of 2 m. 

A nest nearly ready for occupancy, with both male and female bring- 
ing material to it, was a rounded ball placed at the end of a slender leafy 
branch, suspended 2 m above the ground. It was 200 mm or so in 
diameter, made of palm leaves and other fibers, all slender, that pro- 
jected as a rough fringe over the surface. The entrance was low down 
on one side. 

C. O. Handley, Jr., who recorded the wren as abundant, March 20- 
24, 1962, preserved 2 specimens in alcohol. 

It should be emphasized that this wren is distinctly and obviously 
larger than its relative on the mainland, as are the Gould’s Manakin and 
the hummingbird Amazilia handleyi resident on Escudo de Veraguas. 


THRYOTHORUS NIGRICAPILLUS CASTANEUS Lawrence 


Thryothorus castaneus Lawrence, Ann. Lyc. Nat. Hist. New York, vol. 7, June 
1861, p. 321. (Atlantic slope near the Panama Railroad = Lion Hill, Canal 
Zone. ) 


Characters.—Similar to T. n. costaricensis, but paler brown above 
and below; white of the throat usually more extensive, in some reach- 
ing the upper breast; often more heavily barred with black on sides, 
with the barring extending across the lower breast and abdomen. 

A female from the Caribbean slope at the head of Rio Guabal, Cocle, 
taken February 26, 1962, had the iris light mouse brown; maxilla black 
(except the cutting edge); base of mandibular rami dull buffy brown; 
cutting edge of maxilla and rest of mandible greenish neutral gray; 
tarsus and toes fuscous-black. 

A fully grown immature male collected at Gamboa, January 13, 
1960, had the iris light brown; maxilla and a streak on the side of the 
mandible near the tip dull black, the latter changing distally to neutral 
gray, with the base of the mandible and the gape dull honey yellow; 
tarsus and toes dull neutral gray. 


FAMILY TROGLODYTIDAE 89 


Measurements.—Males (14 from the Caribbean slope of Coclé, 
western Colon, and northern Canal Zone), wing 66.2-70.7 (68.9), tail 
49.4-53.7 (51.7), culmen from base 20.3-22.0 (21.0), tarsus 24.9-27.5 
(25.0) tn. 

Females (17 from Coclé, western Colon, and northern Canal Zone), 
wing 63.1-67.0 (64.8), tail 46.3-53.4 (49.2), culmen from base 19.3- 
21.9 (20.2), tarsus 23.4-25.7 (24.7) mm. 

Resident. Locally common on the Caribbean slope from the valley 
of the Rio Calovévora, northwestern Veraguas, through northern Vera- 
guas and northern Coclé (reaching the Pacific slope in these provinces ) 
to the northern Canal Zone (including the middle Chagres Valley), 
and reaching the Pacific slope in the southwestern section. 

In March 1951, I found this wren on the border of the Pacific slope 
near El Valle, Coclé, at 750 m elevation on the base of Cerro La India 
Dormida (male and female collected March 29), and at 600 m at the 
head of the Rio Mata Ahogada (female March 30, male March 31). 
Their presence in this area may be explained by more humid local con- 
ditions in contrast to the drier norm of the rest of the valley, indicated 
by abundant growth of green grass, herbaceous vegetation and vines, 
with occasional areas of new second-growth brush. The actual head of 
the Rio Mata Ahogada, which flows into the Pacific, is immediately 
adjacent to the headwaters of the Rio Indio of the Atlantic drainage, 
the two being separated only by a low, rounded ridge of slight elevation. 
In fieldwork the following year I found this race of the wren common 
inland along the valley of the Rio Indio. In this locality the bird was 
known to the country people as “F1 Guerrero.” The species crosses the 
Continental Divide also in Veraguas at Santa Fé, where it has recently 
been observed by Ridgely and where in 1925 Benson took a specimen 
(AMNH). Eisenmann has also observed it in the fairly humid cano- 
pied second-growth forest on the southwestern section of the Canal 
Zone east of Chorrera. An early report by Salvin (Proc. Zool. Soc. 
London, 1867, p. 134) of a specimen of castaneus labeled “Santiago 
de Veraguas” on the much drier Pacific section of the Isthmus is an 
error as the bird does not occur in that area. Salvin and Godman re- 
peat this locality (Biol. Centr.-Amer. Birds, vol. 1, pt. 4, 1880, p. 88) 
and later authors have also done so. 

srone! (Proc, Acad. Nat.) Sei. Philadelphia; vol: 70) 1918,-p. 271), 
described a nest found by Jewel in the Canal Zone, July 28, 1912, as “a 
loosely built elbow-shaped affair, made almost entirely of a round- 
stemmed grass and lined with finer stems of the same, a few coarser 
stems and reddish-brown vine tendrils on the outside. Loosely placed 


gO BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


in vines, four feet up, by a stream; contained one fresh egg measuring 
.90X.59 in.’ (Equivalent to 2.3<1.5 cm.) The color/et teers 
is not given. 

The notes of E. A. Goldman describe a nest of this species under 
construction near Gatun, Canal Zone, on February 2, 1911. It was 
located about 3.6 m above the ground in dense forest. D. E. Harrower 
(1936 thesis at Cornell University) indicated that the breeding season 
in the Canal Zone ran from late April into August. The season may be 
longer, however, since on November 21, 1957, F. O. Chapelle and G. 
Goldstein found a pair completing a nest near Gatun; an abandoned 
nest was nearby (1m litt. to Eisenmann). 


THRYOTHORUS NIGRICAPILLUS SCHOTTII (Baird) 


Thryophilus schottiu Baird, Rev. Amer. Birds Mus. Smithsonian Institution, vol. 
1, August 1864, p. 123 (in Key), and September 1864, p. 133. (Rio Truando, 
Choco, Colombia. ) 

Thryophilus nigricapillus connectens Chapman, Bull. Amer. Mus. Nat. Hist., vol. 
31, July 23, 1912, p. 157. (Cocal, 5000 ft. elevation, Cauca, Colombia.) 


Characters —Lower surface white, becoming buff on lower abdo- 
men, flanks and undertail coverts, heavily barred with black, in typical 
form this marking extending from the upper foreneck to the undertail 
coverts, in some also over the throat; buff on undersurface darker, 
restricted to flanks, lower abdomen, and undertail coverts. 

A male, taken at Puerto Obaldia, San Blas, March 1, 1963, had the 
iris mouse brown; maxilla black; tip of mandible dull pale olive-green, 
blackish on sides, changing at center to light orange-yellow; gape 
honey yellow; tarsus and toes fuscous-black; claws dark neutral gray. 
An immature female at Pucro, Darién, February 5, 1964, was similar in 
color of iris; maxilla, except cutting edge, fuscous-black; tip of mandi- 
ble dull greenish yellow, changing on side to pale neutral gray; cutting 
edge of maxilla, gape, and basal two-thirds of mandible honey yellow; 
tarsus and toes fuscous-black; claws fuscous. 

Measurements.—Males (10 from Darién), wing 64.0-66.9 (65.7), 
tail 48.0-52.3 (49.7), culmen from base 19.5-22.1 (20.9), tarsus 24.3- 
268) (250, 

Females (10 from Darién), wing 58.2-64.3 (61.9), tail 43.2-49.5 
(46.5), culmen from base 18.6-20.3 (19.6), tarsus 22.3-24.3 (23.4) mm. 

Resident. Common, widely spread through Darién in the upper 
valley of the Rio Tuira, from the Rio Cupe, the Rio Paya, and the Rio 
Pucro to the base of Cerro Tacarcuna; on Cerro Pirre at Cana; on the 
Rio Sambu; along the Pacific Coast from Bahia Pinas to Jaqué, and the 


FAMILY TROGLODYTIDAE QI 


Rio Jaqué, inland to the Rio Imamado; on the Caribbean coast of San 
Blas inland at Armila and Puerto Obaldia. 

In adjacent Colombia this race ranges widely from Acandi and the 
Choco southward. 

Stomachs examined were filled with broken bits of a wide variety of 
insects, and occasionally of a spider, often a fairly large one. One had 
swallowed a small grass seed, apparently a chance occurrence. Six 
collected by Burton (Bull. Brit. Orn. Club, 1975, p. 85) ranged in 
weight from 17.7 to 23.8 g, averaging 20.9 g. 


THRYOTHORUS NIGRICAPILLUS REDITUS (Griscom) 


Thryophilus nigricapillus reditus Griscom, Bull. Mus. Comp. Zool., vol. 72, Janu- 
ary 1932, p. 358. (Permé, San Blas, Panama.) 


Characters.—Similar to T. n. castaneus but somewhat lighter 
colored; white of breast more extensive; sides and abdomen more 
extensively barred with black. 

A fully grown immature female at the Peluca Hydrographic Station 
back of Madden Lake, March 7, 1961, had tarsus and toes black, the 
inner lining of the mouth honey yellow (including the tongue, except 
for a spot of black on the base). An adult female collected near the 
Peluca Hydrographic Station, March 10, 1961, had the tongue black 
except for the tip and two pointed lateral projections at the base, which 
were yellow. 

Measurements.—Males (14 from Cerro Azul, Chiman, Charco del 
Toro, and Cerro Chucanti, in eastern Province of Panama, and Man- 
dinga and Permé, San Blas), wing 67.0-70.5 (68.1), tail 47.5-54.3 
(51.8), culmen from base 19.3-21.9 (20.9), tarsus 24.0-26.5 (25.5) 
mm. 

Females (11 from Cerro Bruja, eastern Colon, Cerro Azul, Chiman, 
Charco del Toro, Cerro Chucanti in eastern Province of Panama; and 
Mandinga, San Blas), wing 63.2-67.7 (65.4), tail 45.0-51.4 (48.5), 
culmen from base 19.0-21.5 (20.1), tarsus 23.1-26.3 (24.7) mm. 

Resident. On the Caribbean side from eastern Colon (Portobelo, 
Cerro Bruja), through San Blas, from Mandinga to Permé. 

A nest seen March 9, 1961, near the Candelaria Hydrographic 
Station was built at the end of a dead branch projecting a few meters 
in the open among the leaves of fairly dense undergrowth on the bank 
of Rio Pequeni. It was an untidy ball of yellowish white fibers as large 
asa coconut. An adult was flushed from the nest, which held 2 nestlings 
with the feather growth barely beginning. 


Q2 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Three collected by Goldman May 25 to 30, 1911, had the stomachs 
filled with fragments of beetles, other miscellaneous insects including 
earwigs, egg cases of roaches, and spiders. 

This population of the wren is definitely an intergrade toward the 
next form, schotti, of eastern Panama and Colombia. Transition be- 
tween castaneus of the Caribbean slope of western Panama east through 
the northern Canal Zone at the western end of its range is fairly abrupt. 
Specimens from near Colon, and northern Canal Zone, are definitely 
castaneus. Those from Portobelo, 30 km east, are reditus. At the 
eastern end the type locality at Permé is barely within the range as 
birds from Armila and Puerto Obaldia, between 15 and 20 km east, are 
placed with schottu. 


THRYOTHORUS SEMIBADIUS Salvin: Riverside Wren, 
Cucarachero Castano Cabecimoreno 


Thryothorus semibadius Salvin, Proc. Zool. Soc. London, November 1870, p. 181. 

(Bugaba, Chiriqui, Panama.) 

Rather small; crown rufous-brown, like back (not black as in nigri- 
capillus) ; rest of undersurface barred narrowly with black and white 
(with 3 black bars on each feather). 

Description.—Length 120-135 mm. Adult (sexes alike) , crown, back, 
scapulars, and upper tail coverts chestnut-brown; wings somewhat 
brownish black, spotted and barred narrowly with light brown on the 
exposed outer webs, the bars extending across both webs on the inner- 
most feathers; lesser coverts black, spotted with white; tail dull black, 
barred and dotted with brownish buff; side of head black, edged widely 
with white, especially on the lores, superciliary line, and cheeks, which 
are largely white; throat white; rest of undersurface, including the 
underwing coverts, white, barred narrowly but heavily with black, the 
white changing to brownish buff on flanks, lower abdomen, and under- 
tail coverts; bend of wing white. 

Juvenile, somewhat duller brown above. 

A male, collected at Corotu, near the coast, northeast of Puerto 
Armuelles, Chiriqui, February 9, 1966, had the iris light reddish brown; 
cutting edge of maxilla and entire mandible neutral gray; rest of 
maxilla black; basal area of inside of mouth black, with anterior half 
of inner surface of maxilla and mandible neutral gray; tarsus, toes, and 
claws neutral gray, with the inner surface of tarsus somewhat paler. 

Measurements.—Males (10 from Chirqui), wing 60.1-66.6 (63.3) 
tail 44.6-50.6 (46.4), culmen from base 19.1-20.6 (19.8), tarsus 22.4- 
25.4 (23.5) mm. 


FAMILY TROGLODYTIDAE 93 


Females (10 from Chiriqui and Costa Rica), wing 60.5-64.7 (62.4), 
tail 41.7-47.5 (44.8), culmen from base 19.2-20.8 (19.8), tarsus 22.4- 
24.5 (23.2) mm. 

Resident. From present information, found locally in the lowlands 
on the Pacific slope of northwestern Chiriqui from the lower Burica 
Peninsula near Puerto Armuelles, to the northern base of the moun- 
tains below and above Concepcion (Bugaba, Divala, Buenavista). 
Thryothorus semibadius is a bird of limited distribution, found in Pan- 
ama in southwestern Chiriqui from the lowlands of the Burica Penin- 
sula near Puerto Armuelles to the base of the mountain area above 
Concepcion. 

My first view of this bird in life came in early March 1960, when a 
few were noted in thickets near the Rio Escarrea, at Buena Vista. In 
February and early March 1966, they were fairly common near Puerto 
Armuelles from the upper Rio San Bartolo, to the Rio Corotu near the 
coast. Like other species of the genus, they ranged in pairs on or near 
the ground in open thickets and in undergrowth in forest, usually near 
water. Ridgely (im litt.) found two pairs in mangroves foraging on the 
roots and in adjacent thickets, at Estero Rico, Chiriqui. Attention is 
drawn to them by their low, chattering calls, and less commonly by 
song, similar to that heard from various races of Thryothorus nigri- 
capillus, but slightly higher pitched in sound. Pairs sing regularly in 
antiphonal concert in which the leader, that 1 assumed to be the male, 
is followed immediately by a lesser voice that I supposed is the female. 
This sequence was clearly demonstrated when I moved quietly along 
a woodland trail, with members of a pair a few feet away, one on the 
right, the other on the left. With the onset of the full dry season at the 
end of February, they sing less often. Because of this silence they are 
seldom noticed in the dense cover that was their shelter. 

In southwestern Costa Rica, Skutch (Pac. Coast Avif. no. 34, 1960, 
pp. 130-137) found it common in the valley of El General, often in 
family groups of 3 or 4 birds. Throughout the year they slept in spe- 
cially built nests in trees or bushes, usually near streams, sometimes 2 or 
3 birds together, possibly a female with grown young, as he believed 
that the mated pairs separated after nesting. The nests made in the 
breeding season were near rapidly flowing streams. They were rounded 
structures approximately 125x150 mm with an entrance chamber and 
an inner compartment. The clutch was two, the eggs being “‘white with 
fine, faint speckles of pale brown that are most numerous in a wreath 
or cap on the thick end.” Three eggs measured 20.6-22.2 by 15.1- 
15.9 mm. The nesting season in Costa Rica extended from late De- 


94 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


cember to September. Skutch found that grown young accompanied 
their parents for several months. On April 20-21, 1968, C. Leck and S. 
Hilty netted birds near the road to El Volcan at ca. 1140 m (confirmed 
by color slide). Eisenmann saw a pair engaged in antiphonal singing at 
Buena Vista (690 m) in a thicket of a boggy area on March 4, 1960. It 
ranges to the west in southwestern Costa Rica from the lowlands to 
about 900 m elevation. 

The close relationship of T. semibadius to the diverse forms of Thry- 
othorus nigricapillus is clearly evident, but the former differs definitely 
and completely from them in always having the crown chestnut-brown 
like the back, and also in the narrow bars on the lower surface which 
are produced by three black bands on each feather, instead of two bands 
as in the races of migricapillus that have this marking. (For other brief 
comment see Wetmore, Smiths. Misc. Coll., vol. 139, no. 2, 1959, p. 17). 
Hellmayr (Cat. Birds Amer., Field Mus. Nat. Hist., pt. 7, 1934, p. 180) 
listed semibadius as a subspecies of nigricapillus, a treatment accepted 
also by Paynter (Check-List Birds World, vol. 9, 1960, p. 407), but 
one that does not seem justified, as there is no indication whatever of an 
approach in the markings described above that separate the two. 


THRYOTHORUS ATROGULARIS Salvin. Black-throated Wren, 
Cucarachero Gargantinegro 


Thryothorus atrogularis Salvin, Proc. Zool. Soc. London, February 1865, p. 580. 
(Tucurrique, Costa Rica.) 


Medium size; deep chestnut-brown; side of head, throat, and upper 
breast black. 

Description.—Length 145-160 mm. Adult (sexes alike); upper sur- 
face, from forehead to upper tail coverts, including scapulars and outer 
webs of wings, chestnut-brown; concealed webs of wings dusky; upper 
tail coverts barred lightly with black; tail black, with indefinite, partly 
concealed brownish bars; side of head to auricular region, chin, fore- 
neck, and upper breast black; superciliary, auricular, and nasal area 
spotted and lined lightly with white. 

Immature, without black on forepart of body, side of head, and un- 
dertail coverts. 

Bill dull black; iris brown, tarsus black. In juvenile, mandible ex- 
cept tip yellowish. 

Measurements.—Males (10 from Bocas del Toro), wing 65.0-68.6 
(66.8), tail 51.4-54.7 (53.2), culmen from base 19.5-22.0 (20.8), tarsus 
21.5-25.5 (24.4) mm. 


FAMILY TROGLODYTIDAE 95 


Females (10 from Bocas del Toro, Costa Rica, and Nicaragua), 
wing 60.0-63.7 (62.2), tail 45.6-50.5 (48.0), culmen from base 19.0- 
20.9 (19.8), tarsus 22.9-24.5 (23.6) mm. 

Resident. Fairly common locally in the lowlands of western Bocas 
del Toro near Almirante Bay; ranging inland to 730 m on the Rio 
Changuena. 

Near Almirante this handsome wren ranges in tangles of low brush 
bordered with stands of coarse grasses. Galindo recorded it inland on 
the lower Rio Changuena, where | was taken September 8, 1961. In 
February and March 1958, they lived locally on the higher ground, 
beyond the borders of cultivation, inland from the wet coastal swamps. 
Attention is drawn to them by their clear musical songs. These are a 
series of modulated notes followed by ringing trills that suggest those 
of Thryothorus fasciatoventris albigularis. In addition to the song, 
they have the usual rattling, chattering notes. The birds ranged low, 
constantly under cover, as 1s usual in the genus. 

The species, here at the lower end of its range, is found north in 
Costa Rica and Nicaragua, through the lowlands of the Caribbean slope. 
The nest and eggs appear to be still unknown. 

Salvin (in Salvin and Godman, Biol. Centr. Amer., Aves, vol. 1, 
1880, p. 91, pl. 6, fig. 4) in his final account of this bird changed the 
spelling of the species name to atrigularis, citing the original spelling, 
atrogularis, in his heading. 


THRYOTHORUS SPADIX (Bangs) Sooty-headed Wren, 
Cucarachero Cabeza Fuliginosa 


Pheugopedius spadix Bangs, Proc. Biol. Soc. Washington, vol. 23, May 4, 1910, 
p. 74. (Naranjito, Rio Dagua, Colombia.) 

Pheugopedius spadix xerampelinus Griscom, Bull. Mus. Comp. Zool., vol. 69, no. 
8, April 1929, p. 182. (Cana, Darién.) 


Medium size; throat and side of head black, the latter area streaked 
with white; chest and back chestnut-brown. 

Description.—Length 140-155 mm. Adult (sexes alike), throat and 
side of head black; a superciliary line and streaks on the cheeks white; 
crown and upper hindneck blackish brown; rest of upper surface, in- 
cluding wings, chestnut-brown; tail black, barred widely with chestnut- 
brown; upper breast dull chestnut-brown; sides, tibia, and undertail 
coverts dull brown, the latter barred with black like the tail; lower breast 
and abdomen dull buff spotted indistinctly with black; undersurface of 
wings buffy white. 

Immature, head, including cheeks and throat, dull brown. 


96 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


Measurements.—Males (10 from Darién, Panama; Antioquia and 
Bolivar, Colombia), wing 64.2-66.9 (65.1), tail 55.1-58.2 (56.9), cul- 
men from base 19.0-20.4 (19.5), tarsus 22.0-25.1 (23.7) mm. 

Females (10 from Darién, Panama; Antioquia and Bolivar, Colom- 
bia), wing 58.2-61.8 (59.0), tail 50.3-54.6 (52.4), culmen from base 
17.8-20.4 (18.8), tarsus 20.7-22.8 (21.7) mm. 

Resident. Found locally in the Subtropical and upper Tropical 
Zone of Darién on Cerro Pirre, Cerro Tacarcuna, and Cerro Quia (780 
m). The population of Panama, named by Griscom, is found to be the 
same as that of western Colombia. 

Goldman recorded 1, taken at 1200 m above Cana, as ranging in un- 
dergrowth along a ridge. The stomachs of 2 that he collected in late 
May and early June were filled with fragments of beetles, hemiptera, 
ants, gryllids, a caterpillar, and spiders. Ridgely (2m ltt.) observed a 
pair foraging in undergrowth at the edge of forest with a mixed flock on 
the slopes of Cerro Quia, Darién, at 540 m, rather lower than most 
other Panama records. The birds were silent as they moved through 
quite open lower undergrowth. In western Colombia, Hilty (Wilson 
Bull., 1974, pp. 479-481) recorded it quite regularly at swarms of the 
small army ant Labidus praedator. There is no record at present of the 
nest and eggs. 

In a considerable series of specimens examined there is no indica- 
tion of intergradation between T. spadix and Thryothorus atrogularis, 
the two groups differing completely in details of the color pattern, nota- 
bly in the crown and upper breast. Hellmayr (Cat. Birds Amer., part 7, 
1934, p. 183) from limited material listed spadix as a geographic race 
of atrogularis, an arrangement followed also by Paynter (in Check-list 
Birds World, vol. 9, 1960, pp. 399-400). The series of both groups 
now available shows no indication of conspecific relationship. Geo- 
graphically the two groups are widely separated. 

S. West (in litt. to Eisenmann, October 1973) reports seeing a wren 
on Cerro Azul, eastern Province of Panama, with a black throat, but 
he failed to determine to which of these species it belonged. 


THRYOTHORUS FASCIATOVENTRIS Lafresnaye: Black-bellied Wren, 
Cucarachero Vientrenegro 
Medium size; foreneck and throat pure white, rest of undersurface 
black, partly barred white, upper surface reddish brown. 
Description.—Length 135-150 mm. Adult (sexes alike), foreneck 
and upper breast pure white, rest of undersurface black, banded nar- 
rowly with buff or with white on lower breast, abdomen and undertail 


FAMILY TROGLODYTIDAE 97 


coverts; central area very faintly lined with the paler markings, or plain 
black; above reddish brown; side of head dull black; a narrow white 
line over loral area and eye. 

Immature, mandible yellow; side of head faintly dusky; light line 
over eye indistinct; foreneck grayish white to gray; rest of undersur- 
face grayish brown to brown, with cross bars faintly indicated or 
absent. 

These are among the more common wrens of forested regions in low- 
land areas, found usually in heavy cover in the denser stands of wood- 
land. Two subspecies are recognized in the Republic. The species 
ranges from southwestern Costa Rica to Colombia, the nominate race 
being described from central Colombia. 


THRYOTHORUS FASCIATOVENTRIS MELANOGASTER Sharpe 


Thryothorus melanogaster Sharpe, Cat. Birds Brit. Mus., vol. 6, “1881” (1882), 
p. 230, pl. 14, fig. 2. (Bugaba, Chiriqui, Panama.) 


Characters.—Slightly darker, more chestnut-brown on upper sur- 
face, and on feathered area of tibia; undersurface with black of breast 
more extensive; averaging slightly larger. 

An adult male, taken at Puerto Armuelles, Chiriqui, February 1, 
1966, had the iris reddish brown; maxilla black; mandible dark bluish 
gray; inner side of tarsus and sides of toes and claws dull neutral gray; 
cuter side and front of tarsus, and tops of the toes dull black. 

Measurements.—Males (10 from Panama and Costa Rica), wing 
65.6-69.6 (66.1), tail 53.2-58.3 (56.7), culmen from base 20.5-23.8 
(22.2), tarsus 23.1-26.2 (24.5) mm. 

Females (9 from Panama and Costa Rica), wing 61.5-66.0 (63.9), 
tail 50.5-55.2 (52.7), culmen from base 19.0-21.9 (20.0, average of 8), 
tarsus 22.5-24.9 (23.5) mm. 

Resident. Found locally in forested areas (not abundant) in the 
lowlands of western Chiriqui; recorded from west of David near 
Divala, and Bugaba, below Concepcion south to Aguacaton, the upper 
Rio San Bartolo and Puerto Armuelles; locally lower on the peninsula 
to Punta Balsa, near Punta Burica. 

As these wrens are primarily inhabitants of thickets in the denser 
cover of forests, they must now be reduced considerably in number 
owing to the extensive clearing in the area from which they are known. 
This western race was known early through specimens collected by 
Arcé, mainly in the Bugaba area (recorded by Salvin, Proc. Zool. Soc. 
London, 1870, pp. 180-181). In the British Museum (Natural His- 


98 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


tory) there are 3 marked with this locality. Several others are labeled 
Chiriqui or “Veragua” without other locality. Griscom’s listing of 
“Veraguas’” in the range probably is due to these specimens, as the bird 
has not been recorded otherwise in that province (Griscom, Bull. Mus. 
Comp. Zool., vol. 78, 1935, p. 358, where the range for Panama is cited 
as “Chiriqui and Veraguas”). Ridgway (U.S. Natl. Mus. Bull. 50, pt. 
3, 1903, p. 532) did mention one locality, Chitra, included in the modern 
Province of Veraguas, but provides no literature basis for this locality 
and there is no specimen in the Smithsonian so labeled. 

Slud in Costa Rica (Bull. Amer. Mus. Nat. Hist., no. 128, 1964, p. 
289) characterizes this form as a “nonforest-inhabiting wren” of the 
densest stream-bordering thickets and wild, tangled brushy growth 
along wooded borders in the humid tropical belt. Its song, “in its slow 
pace and rich dark quality, mellow as that of an oriole,” differs from 
that of other wrens. 


THRYOTHORUS FASCIATOVENTRIS ALBIGULARIS (Sclater) 


Cyphorhinus albigularis P.L. Sclater, Proc. Zool. Soc. London, vol. 23, June 26, 
1855, p. 76, pl. 88. (Isthmus of Panama.) 


Characters.—Slightly lighter brown on upper surface and tibia; black 
of lower parts somewhat less extensive; averaging smaller. 

A male, taken February 6, 1962, at Cafiita, near the Rio Bayano 
(above E] Llano), eastern Province of Panama, had the iris bright 
reddish brown; maxilla black, mandible neutral gray; inside of mouth 
neutral gray, except the tongue which had the distal half translucent so 
that color from beneath appeared through it; inner face of tarsus dark 
neutral gray; rest of tarsus with toes and claws fuscous-black. An- 
other male, collected March 12, 1963, at Armila, eastern San Blas, also 
had the iris reddish brown; maxilla and tip of mandible black; rest of 
mandible neutral gray; tarsus, toes, and claws dusky neutral gray. A 
third of the same sex, collected by R. S. Crossin at Gamboa, August 9, 
1969, was similar. 

Measurements.—Males (10 from Coclé, Canal Zone, and San Blas), 
wing 63.1-70.1 (66.5), tail 49.2-57.1 (53.0), culmen from base 20.0- 
22.2 (20.7), tarsus 23.2-25.9 (24.5) mm. 

Females (10 from Coclé, Colon, Canal Zone, San Blas, and Darién), 
wing 59.7-63.6 (61.3), tail 45.0-50.2 (47.2), culmen from base 19.0- 
21.0 (19.8), tarsus 20.1-24.7 (23.0) mm. 

Resident. Locally fairly common in forested areas, from the Rio 
Indio and its tributaries in western Colon, and the Caribbean slope of 


FAMILY TROGLODYTIDAE 99 


northern Coclé, east on both Pacific and Caribbean slopes in the Canal 
Zone (including Barro Colorado Island where, however, it was last re- 
corded August 22, 1964 [Willis and Eisenmann, Smiths. Cont. Zool. 
291, 1979, p. 24]), through the eastern Province of Panama, including 
the valley of Rio Bayano (Fl Llano, Caftita, Chiman, the Rio Majeé 
(Charco del Toro), and the Tuira Valley (El Real, Pucro); and on the 
Caribbean side through San Blas (Mandinga, Bahia Caledonia, Permé, 
Armila) to the Colombian boundary of northeastern Choco. 

These forest inhabitants, found usually in pairs, at the proper season 
with grown young, range near the ground in low undergrowth, often 
in masses of creepers, low tangles of vines, and dense growths of Heli- 
conia in abandoned banana plantations. Occasionally they were with 
Song Wrens as near companions. The clear, whistled call, wheety-o- 
whee, repeated constantly, usually is heard from birds so well concealed 
in creepers or other cover, from the ground to fairly high in the trees, 
that there may be many minutes without even a glimpse of movement in 
the dense cover that they frequent. Only occasionally will one approach 
almost within arm’s reach, where they utter low calls of a different 
nature and then retreat to more distant cover. 

The song consists of clear hollow whistles, often including or ending 
with a characteristic whee-o-wheet. It is quite different in its more 
varied, constantly changing repertoire, from the rapidly repeated 
phrases of Thryothorus nigricapillus and T. leucotis that often range 
near it in the same forests. 

In stomach examinations the food was found to be the usual variety 
of small insects and spiders, all finely ground. 

In a pair collected by Burton (Bull. Brit. Orn. Club., 1975, p. 85) 
in the Province of Panama the male weighed 28 g and the female 21.4 g. 
Strauch (Bull. Brit. Orn. Club, 1977, p. 64) took a female in the same 
province that weighed 20.7 g. 

One banded at Curundu, Canal Zone, on December 7, 1963, was re- 
captured on September 18, 1967 (Loftin in litt. to Eisenmann). 


THRYOTHORUS RUTILUS HYPERYTHRUS Salvin and Godman: 
Rufous-breasted Wren, Ruisefor del Monte 


Thryothorus hyperythrus Salvin and Godman, Biol. Centr.-Amer., vol. 1, April 
1880, p. 91. (Paraiso station, Panama Railroad, Canal Zone.) 


Small; throat and side of head black, spotted heavily with white; 
undersurface rufous-brown; tail black, barred widely with white. 
Description —Length 115-130 mm. Adult (sexes alike), crown 


100 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


russet-brown, becoming lighter on hind neck; back, including scapulars 
and upper tail coverts, olive-brown; wings somewhat brighter brown; 
tail grayish brown, barred heavily with black, the outer margins of the 
paler bands often white; side of head, including lores and throat, black, 
streaked and spotted narrowly with white; undersurface, including 
sides and legs, tawny-brown, somewhat paler centrally; undertail co- 
verts black, barred with dull grayish white; underwing coverts, includ- 
ing edge of wing, white. 

Adult males, collected at El Copé, Coclé, February 25, 1963, and El 
Volcan, Chiriqui, March 22, 1965, had the iris light reddish brown; 
maxilla and tip of mandible black; rest of mandible somewhat bluish 
neutral gray; tarsus, toes, and claws dark, bluish neutral gray; inside 
of mouth black, this color covering the thickened base of the tongue, 
but not the thin, transparent tip. 

Measurements—Males (10 from Chiriqui, Veraguas, and Canal 
Zone), wing 56.9-60.4 (58.0), tail 47.2-52.1 (48.8), culmen from base 
17.3-18.6 (17.9), tarsus 20.3-23.7 (21.4) mm. 

Females (10 from Chiriqui, Veraguas, Coclé, Los Santos, and Prov- 
ince of Panama), wing 50.6-54.7 (53.6), tail 40.4-46.5 (43.7), culmen 
from base 16.6-18.0 (17.3), tarsus 19.1-20.5 (19.8) mm. 

Resident. Common locally on the Pacific slope from western Chiri- 
qui to the lower Rio Bayano in eastern Province of Panama (not found 
in Darién), including the Azuero Peninsula, and the southern Canal 
Zone; found through the lowlands; ranging upward in hill country at 
El Valle, Coclé, and Cerro Campana, Panama to about 1000 m and in 
Chiriqui to 1600 m (Bajo Mono, Velo) on the Pacific slope of Volcan 
de Chiriqui; recorded occasionally on the Caribbean slope in the Canal 
Zone, especially in the middle Chagres Valley, and at Portobelo, Colon. 

Salvin (Proc. Zool. Soc. London, 1870, p. 181) under the name 
rutilus listed this bird from “‘Calovévora,” and this locality was included 
also in the original description by Salvin and Godman (p. 91), which 
would indicate a locality on the Caribbean slope of Veraguas. 

These are shy birds, widely distributed but not abundant, found in 
pairs along the borders of forest, ranging near the ground, and also 
through tangled vines massed in the lower branches of trees. Their 
tendency to forage higher in low trees makes them somewhat easier to 
see than most wrens. Their presence is usually first made known by 
their constant songs as they creep about under dense cover of leaves. 
Their notes are clear and emphatic, in tone more suggestive of a finch 
or a wood warbler of the O porornis-Seiurus group than of a wren. On 
one occasion, with a companion I followed 1 slowly for sometime 


FAMILY TROGLODYTIDAE IOI 


through its calls, until finally it settled in an area of dense brush. Here 
after watching for 15 minutes, we finally saw it clearly enough to iden- 
tify it. At Sona, Veraguas, in 1953 this wren was called “ruisefor del 
monte.” 

Stomachs of birds collected for specimens were filled with fragments 
of small spiders and a variety of insects, among which were beetles, 
cockroaches, bugs, caterpillars, and ants. One, in addition, held a small 
seed. 

Two males collected by Strauch (Bull. Brit. Orn. Club, 1977, p. 64) 
weighed 15.3 and 15.7 g. 

imiG@osta Riea, Skutch) (Pac. Coast Avif. no: 34/1960, pp. 116-122) 
describes the song and habits in detail. Through the song, given at 
times antiphonally by mated pairs, contact is maintained between in- 
dividuals. He found them building flimsy, covered nests for sleeping, 
which they seemed to occupy alone. In nesting season, pairs built globu- 
lar nests of leaves with an opening in the side. The eggs, two or three, 
were “white with a heavy wreath of brown spots around the thickest 
part, and a sprinkling of the same over the remaining surface. Those 
of one set measured 18.3X13.9, 18.313.9, and 17.9X13.5 millime- 
ters.’ The young have no down at hatching. 

The subspecies hyperythrus ranges from southwestern Costa Rica 
through central Panama, chiefly on the Pacific slope, to the Rio Bayano 
basin. 

The species T. rutilus, as here used, includes other subspecies chiefly 
in hill country and other cleared areas of Colombia, Venezuela, Trini- 
dad, and Tobago. Some authors include in this species the T. maculi- 
pectus group of southeastern Mexico to Costa Rica, and the rather 
similar-looking T. sclatert of Ecuador and Peru (Hellmayr, Cat. Birds 
Amer., pt. 7, 1934, pp. 201-209). Others combine the sclateri group in 
T. rutilus but keep T. maculipectus separate (Paynter, 1960, Check- 
list Birds World, vol. 9, pp. 405-407) or separate T. rutilus but combine 
the sclateri group with the very distant T. maculipectus (Meyer de 
Schauensee, 1966, The Species of Birds of South America, pp. 404- 
405; Howell, Condor, 1957, p. 98). If the species concept of T. rutilus 
is enlarged, the name Speckled Wren is applicable (Eisenmann, im /itt. ) 


TROGLODYTES AEDON Vieillot: House Wren, Cucarachero Comtn 


Troglodytes aédon Vieillot, 1808?, Hist. Nat. Ois. Amer., Sept., vol. 2, p. 52, pl. 
107. (No definite locality mentioned = New York City.) 


Small; short tailed; dark grayish brown, without striking head mark- 
ings; whitish underneath. 


102 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Description—Length 108-115 mm. Adult (sexes alike), brown 
above, with crown slightly grayer; an indistinct superciliary line and 
another back of eye, dull buff; upper tail coverts and rump more ru- 
fescent, the rump with small, concealed white spots and barred nar- 
rowly with black; wing coverts, secondaries, outer webs of primaries, 
and tail dark brown, barred narrowly with dull black; undersurface 
dull white centrally, with sides and flanks brown; undertail coverts dull 
white, mixed with reddish brown and barred with black; scapulars and 
underwing coverts white to pale buff; spotted in some very lightly with 
eray. 

Immature, like adult but without superciliary line; breast and sides 
barred indistinctly and narrowly with dusky. 

The House Wren in Panama is a common bird not only in rural areas, 
but in suburbs and city buildings, where open, warm-weather house 
construction in eaves and corners or holes in brick work often allows it 
entry so that the bird may seem a part of the human family. As a 
friendly neighbor, from its cheerful song, it is popularly known as the 
“ruisenor,” the standard Spanish name for the European Nightingale. 

The species as here treated ranges from Canada and the United 
States through the West Indies, Middle America, and South America 
to Tierra del Fuego and the Falkland Islands. Panama birds belong to 
the T. musculus complex, found from southern Mexico southward 
originally described from Brazil, and often treated as a separate species 
with the name Southern House Wren. 


TROGLODYTES AEDON INQUIETUS Baird 


Troglodytes inquetus Baird, Rev. Amer. Birds, vol. 1, September 1864, pp. 138 
(in key), 143. (Panama Railroad = Atlantic slope, Panama Railroad, Canal 
Zone, Panama.) 


Characters—Undersurface extensively dull white, with grayish 
brown on sides and flanks. 

A male at Aguadulce, Coclé, January 19, 1963, had the iris brown; 
cutting edge of maxilla and base of mandible flesh color; rest of maxilla 
fuscous-black, rest of mandible dark neutral gray; gape honey yellow; 
tarsus, toes, and claws grayish brown. Another male at El Volcan, 
Chiriqui, March 9, 1965, had the iris rather dull reddish brown; base 
of mandible pale brownish white; rest of bill black; tarsus and toes dark 
fuscous; claws dusky neutral gray. In a female, on Isla Cébaco, Vera- 
guas, March 27, 1962, the iris was light, warm brown; maxilla and tip 
of mandible fuscous-black; base of mandible light brownish white; 
gape dull honey yellow; tarsus brownish gray; toes and claws fuscous. 


FAMILY TROGLODYTIDAE 103 


Measurements.—Males (10 from Los Santos, Herrera, Coclé, Prov- 
ince of Panama, Canal Zone, and Colén), wing 51.2-54.2 (52.7), tail 
34.4-37.8 (36.2), culmen from base 15.7-17.8 (16.8), tarsus 18.5-19.8 
(19.3) mm. 

Females (10 from Herrera, Canal Zone, and Darién), wing 51.1- 
54.4 (52.1), tail 34.0-38.0 (35.4), culmen from base 15.4-17.5 (16.0), 
tarsus 18.2-19.5 (18.9) mm. 

Resident. Found locally in the Tropical Zone throughout the Re- 
public, varying locally in abundance; found on Islas Gobernadora and 
Cébaco, Veraguas, and in the Archipiélago de las Perlas, where re- 
ported from Islas Rey, San José, Pedro Gonzalez, Trapiche, Santelmo, 
and Bayoneta). To 2400 m in Chiriqui (trail between Cerro Punta and 
Boquete, Eisenmann, July 13, 1970); 600 m on Cerro Pirre, Darién. 

The House Wren is widely distributed in areas of thickets and fringe 
shrubbery, found in densely forested regions only where clearings have 
been made. Normally the birds are in pairs, or for brief periods, in 
family groups. They come into cities and towns, and about country 
homes are regular attendants, searching corners, thatched roofs where 
present, and any other concealing cover for their insect food. As they 
move about, males sing constantly their rapid, bubbling songs, joined 
regularly by the wheezy notes of the females. Nests of soft vegetable 
fibers, often with a feather lining, are hidden in cavities, usually with 
narrowed entrances so that eggs and young are completely hidden. 
Construction is by both members of the pair, incubation by the female 
alone, but she is fed regularly by the male, who also aids in care of the 
young. In the breeding season from March to July (sometimes later) 
three broods in rapid succession are usual, with young of the first family 
frequently aiding in bringing food to those that follow. 

The eggs are white, marked rather heavily with pale to dark brown; 
usually there are 3 or 4, rarely 5 toa set. They range in size from 16.7- 
19.4 12.7-13.9 mm, with an average of 17.8X13.4 mm. 

Males sing more or less regularly through the year except when molt- 
ing, with young males beginning rambling songs when 2 months old or 
less. Full details on this species in Central America are given by Skutch 
(Condor, 1953, pp. 124-149; Naturalist in Costa Rica, 1971, pp. 22-24). 

Adult birds of this subspecies are fairly uniform in color throughout 
the Isthmus, those from the base of the Burica Peninsula being similar 
to those of eastern Panama. The only variation noted is in 1 male col- 
lected in the uplands of northwestern Chiriqui within 15 km of the 
Costa Rican boundary. This bird is intermediate between T.a. inter- 
medius and the Panamanian inquietus, the entire upper surface and the 


104 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


sides being dark like intermedius but with the central undersurface 
nearly clear white as in inquietus. The wrens, not common in this area, 
should be examined from within a kilometer or so of the boundary, as 
they may prove to be the subspecies intermedius. Griscom ascribed 
birds from extreme northwestern Panama in Bocas del Toro (Almi- 
rante, Chiriquito) to intermedius, but they are here included in 
inquietus. 

Birds weighed by Burton (Bull. Brit. Orn. Club, 1975, p. 85) and 
Strauch (Bull. Brit. Orn. Club, 1977, p. 64) weighed from 14.0 to 
LZ 

Nests with eggs or young have been reported in Panama in every 
month from January through October, according to Eisenmann, but 
egg laying may occasionally occur in the other two months. On Feb- 
ruary 5, 1960, he saw a stub-tailed fledgling accompanying its parent. 

The bubbling, presumably territorial, song strongly resembles that 
of T. a. aedon of eastern United States, but there is another musical 
vocalization commonly given even during the months November to 
February, when the bubbling song is infrequently heard. Eisenmann 
has never heard nominate aedon give a call resembling it. The vocaliza- 
tion is a loud, clear, rapidly repeated chwee-chwee-chwee and so on, all 
on one pitch, sometimes more like chew-chew-chew. Occasionally, the 
same bird will shift from this vocalization to the irregular, much more 
varied bubbling song. Eisenmann has also heard in Panama scolding 
and chattering like churkat and churr-cheéchee. 

On Barro Colorado Island this species, when present, comes to pick 
off window screens or the pavements insects that were killed when at- 
tracted to lights. Eisenmann has seen House Wrens walking, as well as 
hopping, on the ground without any clumsiness. 

Although recorded from Bocas del Toro, it is, or was, apparently 
rather rare there, for on three trips to the Almirante Bay area (the last 
in 1965) Eisenmann failed to observe it. 


TROGLODYTES AEDON CARYCHROUS Wetmore 


Troglodytes aedon carychrous Wetmore, Smiths. Misc. Coll., vol. 134, no. 9, July 

8, 1957, p. 76. (Isla Coiba, Panama.) 

Characters.—Similar to T. a. inquietus but much brighter brown 
throughout; upper surface definitely deeper brown; underneath with 
sides and flanks darker brown, this color in some covering the entire 
surface. Others are whiter on throat and center of breast. 

In its definitely brighter coloration this bird of Coiba is distinct from 


FAMILY TROGLODYTIDAE TO5 


the race inquietus of Panama, somewhat less so from intermedius found 
in Costa Rica. Three juveniles taken January 26, 1956, as they were 
about to leave the nest, differ from those of similar age of inquietus. 

Measurements.—Males (11 from Isla Coiba), wing 51.4-53.5 (52.2), 
tail 34.0-37.1 (35.5), culmen from base 16.9-19.1 (17.8), tarsus 18.6- 
20.5 (19.4) mm. 

Females (2 from Isla Coiba), 49.0-50.1 (49.5), tail 31.5-33.2 (32.3), 
culmen from base 17.3-17.4 (17.3), tarsus 19.4-19.9 (19.6) mm. 

Resident. Coiba Island, prior to modern development, was heavily 
forested almost throughout; in 1976 it was still 75 percent forested 
(Ridgely in litt.) The House Wren perforce was definitely a forest in- 
habitant, far more so than in other areas of Panama, except in part in 
the Pearl Islands. In Coiba, it was common in the lower growths near 
the beaches and the swampy woodlands bordering the mangroves in 
those areas. But also I found it regularly in undergrowth through the 
high interior forests. There the birds were seen in the main around fal- 
len trees or where undergrowth was covered with masses of vines. Only 
occasionally did they appear to range higher in the trees. In these lo- 
cations they remained under cover, while around the main buildings of 
the penal colony they were less timid. In their usual activities their 
songs, heard daily, were a constant pleasure. Ridgely (im litt.) finds 
the song of this race richer and more melodic than that of mainland 
forms. 


TROGLODYTES SOLSTITIALIS Sclater: Mountain Wren, 
Cucarachero de Sierra 


Troglodytes solstitialis P.L. Sclater, Proc. Zool. Soc. London, vol. 26, 1858, p. 550. 
(Matos and Pinipi, near Riobamba, Ecuador.) 


Small; smaller than Troglodytes aedon, with side of head and su- 
perciliary line definitely brighter brown. 

Description.—Length 92-100 mm. Dull reddish brown above, and 
in the line behind the eye; wings and tail narrowly barred with black; 
undersurface centrally white; sides and flanks buffy brown; undertail 
coverts buffy brown, barred narrowly with dusky. 

Two forms are recognized in the higher mountain areas of Panama, 
one in Chiriqui, the other in Darién. Few specimens have been col- 
lected. 

The Mountain Wrens from Costa Rica and Panama are sometimes 
given species rank as T. ochraceus, Ochraceus Wren. Here they are 
treated as subspecies of the South American T. solstitialis group found 
in the high Santa Marta mountains of Colombia and the Andes to 


106 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Venezuela, Ecuador, Peru, Bolivia, and northwestern Argentina. 
Paynter (Check-List of Birds of the World, vol. 9, 1960, pp. 427-429) 
also includes the T. rufociliatus group of southern Mexico (Chiapas), 
Guatemala, Honduras, and El Salvador in T. solstitialis. 


TROGLODYTES SOLSTITIALIS LIGEA Bangs 


Troglodytes solstitialis ligea Bangs, Proc. New England Zool. Club, vol. 4, March 
19, 1908. p. 29. (Volcan de Chiriqui, 1220 m above Boquete, Chiriqui.) 

Troglodytes ochraceus remotus Griscom, Amer. Mus. Nov. 141, 1924, p. 5. (Cerro 
Flores, alt. 6000 ft., Chiriqui.) 


Characters.—Brighter, more reddish brown on dorsal surface, in 
the superciliary line on side of head, and on the flanks; slightly larger 
than T. s. festinus. 

An adult female, taken on Silla de Cerro Pando, beyond El Volcan, 
Chiriqui, March 13, 1954, had the maxilla dark neutral gray, except the 
cutting edge which was marguerite yellow for the basal third, shading 
anteriorly to mouse brown; basal area of cutting edge in mandible also 
marguerite yellow; mandibular rami light yellowish, with a line of neu- 
tral gray at tip and along the side; iris brown; tarsus and toes mouse 
brown. 

Measurements.—Males (10 from Chiriqui), wing 47.1-49.9 (48.6), 
tail 28.6-36.2 (29.6), culmen from base 14.5-16.2 (15.2, average of 9), 
tarsus 16.5-18.0 (17.4) mm. 

Females (8 from Chiriqui), wing 43.6-48.6 (45.1), tail 27.3-33.1 
(28.8), culmen from base 14.6-15.5 (15.0), tarsus 16.0-17.9 (16.8) 
mm. 

Resident. Upper levels of Volcan de Chiriqui, 900 to 2500 m, west- 
ern Chiriqui. 

This small wren, compact and rotund in form, is a forest species, 
secretive in habit. I found them near and on the ground, in dense 
tangles, rarely amid fallen tree trunks in small, recently made clearings. 
Once only, in dense forest, did I see one range 15 m above the ground 
in a mass of tangled vines near the top of a dead stub. Those seen were 
silent. 

In the Monniche collection, Blake (Fieldiana: Zool., vol. 36, no. 5, 
1958, p. 547) recorded a small series taken on the slopes of the volcano 
above Boquete, between 1580 and 2500 m elevation. 

Griscom described remotus from a single female taken in the moun- 
tains back of Remedios, Chiriqui. Some form of this species (ochra- 
ceus group) definitely occurs in the mountains of eastern Chiriqui (Rio 
Chiriqui, 600-1140 m, near the Fortuna Dam site, where between Feb- 


FAMILY TROGLODYTIDAE 107 


ruary 26 and March 5, 1976, Ridgely found it common at all elevations) , 
Veraguas (above Santa Fé along Cerro Tute ridge, 1050 m, January 5, 
1974), and probably even in western Province of Panama (at the very 
top of Cerro Campana, ca. 1000 m, seen by N. G. Smith on September 
7, 1968). 

Eisenmann has frequently seen it in humid montane forest from the 
lake near El Volcan, ca. 1200 m, to well above Cerro Punta, close to 
2400 m, and also on the Boquete side above Finca Lerida. They gen- 
erally glean over vines and similar epiphytic vegetation on forest trees, 
‘usually in pairs, creeping vertically along hanging lianas, but occas- 
ionally even up tree trunks. Ridgely (1m litt.) notes that this species 
is a rather frequent member of mixed flocks including warblers and 
tanagers. 

Eisenmann noted a semi-musical trilled tswee-tswee seeerrrr, also a 
seecerrrrr alone, lasting about one second, sometimes drier as whirrrrr, 
and a short, nonmusical srr or srrrrr or trrrr, which is perhaps the call. 

On April 26, 1961, on Martyn’s Finca in the still forested, but logged 
over area, Kisenmann saw pairs actively singing and displaying. One 
bird landed on a vertical trunk and sang with shivering wings; another 
carried nesting material. Still another pair had a nest or were building 
in a hollow of a fallen trunk or nearby, for they would not leave the 
vicinity even on close approach. 

Skutch (Pac. Coast. Avif., 34, 1960, pp. 166-167) describes behavior 
and nesting in Costa Rica (ochraceus), and states nests are placed in 
cavities on broken ends of decaying branches or trees and that nesting 
must begin by April. He found another pair building in late May. 
Nests of solstitialis in Ecuador were in different sites. 


TROGLODYTES SOLSTITIALIS FESTINUS Nelson 


Troglodytes festinus Nelson, Smiths. Misc. Coll., vol. 60, no. 3, September 24 
(September 27), 1912>p.22) (Cerro Pirre 1580 m; Darien.) 


Characters.—Similar to T. s. ligea, but slightly smaller; somewhat 
darker on upper surface and on sides and flanks. 

Measurements.—Males (3 from Darién), wing 43.9-44.5 (44.1), 
tail 27.2-30.4 (29.1), culmen from base 14.9-16.5 (15.7), tarsus 17.1- 
7-3 (17.2) mom. 

Female (1 from Darién), wing 39.4, tail 28.0, culmen from base 12.6, 
tarsus 16.0 mm. 

Resident. Recorded from the upper levels of the mountains of east- 
ern Darien; Cerro Pirre, 1580 m at the head of Rio Limon; and Cerro 


108 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


Mali, at 1460 m (6 km west of the summit), on the base of Tacarcuna. 

The birds of the Darién mountains differ only slightly from those of 
Chiriqui in size, and in darker coloration on the upper surface and sides. 

The race festinus was described from a single adult male, collected 
by E. A. Goldman in the forest near the head of Rio Limon. It is still 
the only specimen known from Cerro Pirre. From June 4 to 9, 1963, 
Galindo secured 2 males and a female in the forest adjacent to his camp 
near the summit of Cerro Mali, on the side of Cerro Tacarcuna. 

The 3 birds from the Tacarcuna massif are slightly darker on the 
back and crown of the head than the single male from Pirre; with more 
material they may prove to be subspecifically distinct. From the few 
now available it is noted that the slight differences seen are, in the main, 
equivalent to the individual variation noted in an extensive series of the 
race ligea from Chiriqui. 


TROGLODYTES BROWNI BROWNI Bangs: Timberline Wren, 
Cucarachero de Volcan 


Troglodytes browni Bangs, Proc. New England Zool. Club, vol. 3, January 30, 
1902, p. 53. (3000 m elevation, Volcan de Chiriqui.) 


Small, short-tailed; reddish brown above, paler on flanks and under- 
tail coverts; with a prominent white superciliary, and a brown line (in 
color like the crown) through the eye. 

Description.—Length 95-100 mm. Adult (sexes alike), above some- 
what dull reddish brown, slightly darker on crown, brighter on rump; a 
prominent white superciliary line, and a brown line (like crown) 
through eye; wings and tail dusky, barred with reddish brown; lesser 
wing coverts flecked with white; outer primaries edged with white; 
sides of neck dull white edged with dusky brown; undersurface chin to 
abdomen, dull white edged faintly with dusky; flanks and undertail 
coverts dull russet-brown, the undercoverts flecked with white; edge of 
wing and underwing coverts dull white. 

Immature, superciliary dull white; feathers of undersurface mar- 
gined with dusky; white markings on primaries more extensive. 

Measurements.—Males (8 from Chiriqui), wing 48.8-53.1 (51.3), 
tail 28.9-33.2 (31.5), culmen from base 14.1-16.0 (15.0), tarsus 21.9- 
25./ (25/6) aa: 

Females (8 from Chiriqui), wing 47.6-54.1 (50.3), tail 28.4-32.2 
(30.3), culmen from base 14.0-16.1 (14.8, average of 7), tarsus 21.8- 
24.0 (22.4) mm. 


Resident. Common near timberline across the summit of Volcan de 


FAMILY TROGLODYTIDAE 109 


Chiriqui. It is found also on the highest mountain peaks of Costa Rica. 
This wren was described by Bangs from a series of 11 specimens col- 
lected by W. W. Brown, Jr., in May and June, 1901. The collector 
found this species ‘wholly confined to the cane brakes on top of the 
Volcan de Chiriqui ... It lived much after the manner of a marsh wren, 
and its song and notes were wholly unlike those of any wren” known to 
him. In the American Museum of Natural History there are 2 females 
collected by H. J. Watson, October 1 and 4, 1904. On the labels the 
collector noted colors of the soft parts as follows: “Iris violet, feet 
brown, bill black.” From the Monniche collection Blake recorded 2 
females July 8 (year not indicated) from the summit of the volcano 
above Copete, and 2 males and 2 females July 12, at about 4200 m 
elevation. 

Little is recorded of these birds. Worth, in a brief note (Bird-Lore, 
vol. 41, 1939, p. 282), wrote that they live in bamboo thickets, and that 
the song “is unlike that of any House Wren I have heard. It is not a 
finished entity, like a House Wren’s performance, but only a cyclically 
repeated phrase without musical beginning or end; it could... be as 
short or prolonged as desired. The scolding note . . . however, bears 
some resemblance to the House Wren’s complaining, but it lacks the to- 
and-fro rasping quality of the House Wren, being a single utterance of 
uniform timbre.” 

In Costa Rica, Slud (Bull. Amer. Mus. Nat. Hist., vol. 128, 1964, p. 
291) found closely allied races inhabiting “the bushy scrub and impen- 
etrable canelike bamboos that border the high-altitude forests, at breaks 
and above the limit of trees. Though usually met singly or in twos, a 
number of individuals may be scattered over an area. This stub-tailed, 
white-winged little wren is active in the dense low cover; it may be 
silent but it is not shy. The behavior and the thickety habitat are typi- 
cal of a wren. Its patterned song is thin, not very clear, rather mea- 
sured, wrenlike.” C. Hartshorne, regarding the song at Cerro de la 
Muerte, Costa Rica (3,090 m), says (im litt. to Eisenmann) it is as 
long or longer than that of the Winter Wren, but not quite as loud or as 
musical. It “consists of two slightly contrasting phrases, repeated fast 
over and over, something like “bussell-bissell, bussell-bissell,’ etc. for as 
many as ten or more seconds. This is a crude description except for 
the idea of the slight contrast and the numerous quick repetitions.” 

The nest and eggs are not known. 

Oberholser (Proc. U. S. Nat. Mus., vol. 27, 1904, p. 198) proposed 
for it the separate genus Thryorchilus on the basis that it has only 10 
rectrices instead of the 12 found in Troglodytes. 


110 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


HENICORHINA LEUCOSTICTA (Cabanis): White-breasted Wood-Wren, 
Cucarachero Pechiblanco 


FicureE 11 


Cyphorhinus leucostictus Cabanis, Arch. Naturg., Jahrg. 13 Bd. 1, Heft 2, 1847, 
p. 206. (Guiana and Mexico (Papantla) = British Guiana; designated by 
Berlepsch and Hartert, Nov. Zool., vol. 1X, April 1902, p. 5.) 


Small; tail short; foreneck and breast white; side of head black, 
varied with white lines; a white line over eye. 

Description.—Length 92-105 mm. Back, wings, and tail chestnut- 
brown; crown dark, varying from grayish brown to black; throat to 
upper abdomen pure white, without markings; flanks tawny. 


Ficure 11.—White-breasted Wood-wren, Cucarachero Pechiblanco, Henicorhina 
leucosticta. 


Juvenile, throat white; foreneck and breast dull gray; colors and 
markings otherwise as in adult. 

A forest inhabitant, widely distributed. This common species of the 
lowland forests may be recognized by the pure white of the foreneck 
and breast. The two subspecies accepted here from Panama are closely 
similar in size, differing in depth of color in the darker markings of the 
crown. 

The species ranges from eastern Mexico, Belize, the Caribbean slope 
of Central America (extending to the Pacific slope in Costa Rica and 
Panama), through Colombia, Ecuador, and Peru, southern Venezuela, 
the Guianas, and the upper Rio Negro region of northern Brazil. 


FAMILY TROGLODYTIDAE EVE 


HENICORHINA LEUCOSTICTA PITTIERI Cherrie 


Henicorhina Pittieri Cherrie, Anal. Inst. Fis.-Geogr. Mus. Nac. Costa Rica, vol. 
IV, 1891 (1893), p. 134. (Boruca, Puntarenas, Costa Rica.) 

Henicorhina prostheleuca tropaea Bangs and Peters, Bull. Mus. Comp. Zool., vol. 
Ixvii, no. 15, January 1927, p. 480. (La Vijagua, Alajuela, Costa Rica.) 


Characters.—Closely similar to H. |. darienensis, but with crown 
median area washed with rufescent brown (variable in depth of color). 

A male collected at the head of the Rio Guabal, on the Caribbean slope 
of Coclé, February 28, 1961, had the iris dark mouse brown; bill black; 
tarsus, toes, and claws dusky neutral gray. 

Measurements.—Males (10 from the Pacific slope, from south- 
western Costa Rica, and western Chiriqui, eastward through the 
Canal Zone), wing 52.5-58.6 (55.0), tail 23.0-29.1 (26.5), culmen from 
base 17.0-18.9 (17.8), tarsus 22.5-24.7 (23.6) mm. 

Females (10 from western Costa Rica, Chiriqui, and Bocas del Toro 
to the Canal Zone, and Chepo, eastern Province of Panama), wing 
eo o2.1), tail 23'022710' (25.5); ‘culmen’ from base 15.6-17.7 
(16.6), tarsus 21.6-25.5 (22.6) mm. 

Resident. Common in forests to 1300 m, less so to 1900 m; on 
Pacific and Atlantic slopes from the Costa Rican boundary through 
western Chiriqui and western Bocas del Toro (Almirante, Cricamola, 
to 750 m above Rio Changuena). Found also on the Caribbean slope of 
Coclé (upper Rio Coclé del Norte, Rio Calovévora), and western Colon 
(on the Rio Indio); on the Pacific side, Veraguas (Chitra, Santa Fé), 
Coclé (El Valle), on Cerro Campana, western Province of Panama 
and the Cerro Azul-Cerro Jefe area, the hills north of Chepo (Zanja 
Limon), eastern Province of Panama. In the Canal Zone it is found in 
woodlands at Fort Clayton, Madden Forest Preserve, Pipeline Road 
area, and along the Achiote Road. Birds collected by Ridgely and Ga- 
lindo in the lowlands and hills above the Rio Bayano and its tributaries 
have not been subspecifically identified. 

Northward in Central America, this form ranges on the Caribbean 
slope through forest areas in Costa Rica, Nicaragua, Honduras, and 
Guatemala. With the series of specimens now available the supposed 
subspecies tropaea named by Bangs and Peters from La Vijagua in 
the Caribbean drainage of northern Costa Rica, with a range extending 
east to Bocas del Toro and northward to Guatemala, is not separable 
from pittveri. 

On the mountain slopes in western Chiriqui I found these wrens com- 
mon in pairs in undergrowth through the forest. As they moved about 
in dense cover, usually near the ground, they were recorded mainly 


I12 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


from their clear ringing songs, and their chattering calls. In higher 
elevations they often frequented bracken and other tangled growth on 
steep slopes in treeless pasturelands. Brown collected it in the Boquete 
district at about 1800 m, but it is rare at such high elevations. In the 
lowlands of Panama it favors damp, canopied forest, especially in hilly 
areas and about ravines. 

The song is a loud, beautifully rich series of whistles, usually rather 
short, mostly of three or four phrases, repeated many times and then 
varied. The most characteristic song is churry-churry-cheeer (inter- 
preted mnemonically by F. O. Chapelle as “pretty-pretty-bird”), and 
variations; Eisenmann has also heard whee-tew-tew and choodweeoo- 
chooweréh chwee-tew-tew. One vocalization suggested a slightly aber- 
rant song of the Green Shrike-Vireo syllabized as teea-teea-teeoo. Calls 
include a peculiar bweeer or beeer, which reminded Ridgely of a note 
of the Swainson’s Thrush. E. S. Morton has made a tape recording 
(Cornell Laboratory of Ornithology) of a hoarse, throaty, rather frog- 
like hreer or hreéyee (syllabized by Eisenmann from the recording). 
Usually the bird sings from a concealed position, very low in a tangle 
or thicket, but occasionally in the dense vegetation of a tree. 

This species is not infrequently attracted to army-ant swarms, and 
may also forage below wandering bands of antbirds. The small tail is 
often carried cocked up. 

Skutch (Pac. Coast Avif. no. 34, 1960, pp. 140-143) records that 
they build a rather flimsy nest for sleeping, a rounded ball of partly 
weathered leaves, with an opening at one end, placed in forest under- 
growth elevated 1 or 2 m, sometimes more, above the ground. Adults 
may be accompanied by a single young bird, but otherwise sleep alone. 
Nesting structures for eggs and young were more substantial. The 
eggs, two in number, were clear white, with measurements of 14.3-15.1 
x 20.6-23.0. 

In another account of nest and eggs (identified as those of this race 
but listed under the synonym tropaea), Huber (Proc. Acad. Nat. Sci. 
Philadelphia, vol. 84, 1932, p. 238), on May 19, 1922, in forest back of 
the Eden mine, northeastern Nicaragua, near the headwaters of the 
Rio Bambana, flushed one of these wrens “from a nest containing two 
fresh eggs. The nest, about eighteen inches above the ground, was 
placed in a crevice in a fallen log. This old moss-covered log lay close 
to a trail in the heavy forest and the jar from my weight . . . flushed the 
bird. The nest, a carefully made, bulky, globular-shaped affair com- 
posed of fern stems, plant fibers, and moss, is lined with fine grasses. 
The entrance is from the side. The outside diameters are 190 X 130 mm. 


FAMILY TROGLODYTIDAE LES 


The eggs are white with a very few light brown specks. They measure 
20.0 x 14.5, 20.1 14.1 mm.” 

The race pittieri was named by Cherrie in honor of Professor Henri 
Pittier, resident at that time in San José, Costa Rica, and active in 
natural history studies in that country. 

A male taken by Strauch (Bull. Brit. Orn. Club, 1977, p. 64) at 
Cerro Campana weighed 16.1 g. 

Morton (in Temple, Endangered Birds, 1978, pp. 379-384) describes 
a reintroduction of this species to Barro Colorado Island, from which 
it had disappeared before 1960. 


HENICORHINA LEUCOSTICTA DARIENENSIS Hellmayr 


Henicorhina leucosticta darienensis Hellmayr, Anz. Ornith. Ges. Bayern, Bd. I, 
no. 4, March 1921, p. 25. (“Tacarcuna dstliches Panama” = former Tacarcuna 
Village site, west base of Cerro Mali, 600 m elevation, Cerro Tacarcuna, 
Darién. ) 

Characters.—Crown and upper hindneck plain deep black; brown 
of back, wings, flanks, and undertail coverts somewhat lighter and 
brighter. 

Measurements——Males (10 from San Blas, eastern Province of 
Panama, and Darién), wing 53.4-55.5 (54.4), tail 22.5-25.5 (24.3), 
culmen from base 16.8-18.6 (17.6), tarsus 21.3-24.7 (22.4) mm. 

Females (10 from eastern Province of Panama, San Blas, and 
Darién), wing 50.0-53.4 (51.6), tail 21.2-24.0 (23.0), culmen from 
base 15.6-16.5 (15.9), tarsus 19.3-22.3 (21.1) mm. 

Resident. Common in forested areas from western Comarca de 
San Blas (Mandinga) and Chiman (hill country on Rio Chiman), east- 
ern Province of Panama, east to the Colombian boundary, on both 
Caribbean and Pacific slopes; in Darién from near sea level to 1000 m 
on Cerro Pirre, 1450 m on Cerro Tacarcuna, and 930 (the summit) of 
Cerro Quia. This race also occurs in adjacent northwestern Colombia, 
on both slopes, to the Sint Valley and the Baudo Mountains. 

While widely distributed in forest areas, in general this subspecies is 
less abundant than the race pittieri of western Panama. A male from 
the Rio Chiman at the mouth of Rio Corotut is typical of this race in 
black color of the crown. Found as usual in pairs, they were especially 
common on the upper Rio Jaqué in Darién, and elsewhere in the in- 
terior. None were recorded near the coast at Jaqué. Like the other 
subspecies these are birds of the forest, living in the shelter of dark 
shadows. 


II4 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


The food, from examination of stomach contents, includes the usual 
variety of small insects—ants, small beetles, caterpillars, earwigs, 
roaches and their eggs, and small spiders. One collected by Burton 
(Bull. Brit. Orn. Club, 1975, p. 85) at Cerro Pirre weighed a2 


HENICORHINA LEUCOPHRYS (Tschudi): Gray-breasted Wood-Wren, 
Cucarachero Pechigris 

Small, with short tail, similar to Henicorhina leucosticta, but breast 
gray and tail longer. 

Description.—Length 103-115 m. Back, wings, and tail chestnut- 
brown; crown sooty black; side of head black, lined prominently with 
white, with a long white superciliary line; tail dull brown, barred more 
or less with black; wings like back, indistinctly barred with black; an- 
terior undersurface gray, with throat grayish white, plain in the nomi- 
nate race, lined heavily with gray in subspecies collina; flanks and un- 
dertail coverts reddish brown. 

Juvenile, darker above with the crown dull black; a prominent white 
superciliary; side of head dull black, with the white markings, com- 
pared to the White-breasted Wood-Wren, somewhat reduced; throat 
for a limited area dull white; foreneck and upper breast dark gray; rest 
of undersurface dull brownish black. 

A forest inhabitant, widely distributed in highland areas. Common 
on wooded slopes, only in the mountains. Two subspecies are found on 
the Isthmus, one in the west, the other, less common, from Cerro Azul 
eastward. 

The species ranges from the highlands and mountain slopes of east- 
ern Mexico, through Central America to Colombia, Venezuela, and in 
the Andes (chiefly at subtropical altitudes) of Ecuador and Peru to 
Bolivia. 


HENICORHINA LEUCOPHRYS COLLINA Bangs 


Henicorhina collina Bangs, Proc. New England Zool. Club, vol. 3, Jan. 30, 1902, 
p. 55. (Boquete, Chiriqui, 1800 m alt.) 


Characters.—Throat dull grayish white, heavily streaked with dark 
gray; otherwise like H. 1. leucophrys. 

M easurements—Males (10 from Chiriqui and Veraguas), wing 54.6- 
59.2 (57.0), tail 30.5-34.3 (32.0), culmen from base 17.1-19.0 (18.1), 
tarsus 24.2-25.6 (25.0) mm. 

Females (10 from Chiriqui), wing 52.3-55.6 (53.6), tail 27.1-29.0 


FAMILY TROGLODYTIDAE ; I15 


(27.9), culmen from base 16.0-17.5 (16.5), tarsus 22.5-25.0 (23.9) 
mm. 

Resident. Common on the Pacific slope of the central mountain 
range in Chiriqui and Veraguas, from near the Costa Rican boundary 
to Cerro Campana, western Province of Panama, ranging from 900 to 
2900 m. Recorded also from the upper levels of the Caribbean side of 
the Divide. Found at 900 m elevation on Cerro Campana, western Prov- 
ince of Panama. Beyond Panama this race occurs only in the highlands 
of Costa Rica. 

This species is widely distributed over the mountain slopes, mainly 
forest, but also where steep open slopes are grown heavily with bracken. 
The main distribution is on the Pacific drainage, but Blake (Feld: 
Zool., vol. 36, no. 5, 1958, p. 547) records specimens from the Holcomb 
Trail above Boquete on the Caribbean side of the high divide in Bocas 
del Toro. It ranges eastward along the Pacific slopes to near Santa Fé 
and Chitra, Veraguas, an area in which it also doubtless crosses the 
Divide, as it is recorded from Calovévora. Ridgely (im litt.) found it 
very common in late February and early March 1976 in forest under- 
growth at the Fortuna Dam site in central Chiriqui (900-1500 m), 
where no H. leucosticta were present. In one day he netted 13 indi- 
viduals. 

It is found in pairs, their presence known by the steady songs of the 
males, in which the females regularly join. An intruder may be greeted 
with a harsh scolding note, repeated quickly several times. While these 
birds remain in areas where they have cover, often they seem more con- 
fiding than some other forest wrens as they may be seen without too 
great difficulty. 

Carriker (Ann. Carnegie Mus., vol. 6, 1910, p. 762) described a nest 
found on Irazu, Costa Rica, April 17, 1902, that held two plain white 
eggs without markings. These measured 21.5xX15 and 22X15 mm. 
Skutch, in a detailed account (Pac. Coast Avif., no. 34, 1960, pp. 146- 
154), records that the pair sleep together in rounded, covered nests with 
the entrance underneath. Adults may be accompanied in the nest by 
their young for a period after these are grown. Skutch describes the 
two eggs in a set as plain white with measurements of both in one nest 
as 22.2 X15.5 mm. 

A male collected by Strauch (Bull. Brit. Orn. Club, 1977, p. 64) at 
Cerro Campana weighed 18.7 g. 

This species is very common in the highlands of western Chiriqui, 
especially above 1500 m on both the east and west slopes of the volcanic 


116 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


massif, and is often seen or heard in this area along roadside thickets 
and hedgerows separating open fields, at a distance from surviving for- 
est. In this highland region, Eisenmann found it the commonest wren, 
having adjusted to clearing and cultivation, and calling attention to itself 
by persistent singing, even through it skulks in vegetation, generally 
from a few centimeters to 1 or 1.3 m from the ground. Notably abun- 
dant in the Cerro Punta area, now largely devoted to vegetable crops, it 
frequents the shrubbery at field boundaries (as well as forest borders 
and interiors). In this region it occurs below El Volcan at least down 
to 1110 m along the highway, and about the Volcan lakes. It occurs also 
at the summit of the trail to Boquete, and seems to be everywhere in the 
outskirts of Boquete, even in the rather scantily vegetated El Salto. On 
Cerro Campana, where sympatric with H. leucosticta, it is much less 
numerous than that species and is confined to humid forest about the 
summit. 

This species less frequently carries its relatively longer tail cocked 
up than does the White-breasted Wood-Wren, and its rufescent lower 
underparts are conspicuous in the field. 

The song, although musical and loud, is not of as rich or mellow a 
quality as that of its congener, but usually consists of many more 
syllables, and lasts longer before it 1s repeated. One bird seen by Eisen- 
mann on September 20, 1958, gave three different songs, all loud, 
rollicking whistles of similar timbre. He noted the following songs: too- 
tee-ooweét, weétee, weétoo, also weeloo-weéchee, too-loo-wheew, and 
cheerooeecheé-cheeweécheerooweéchee, with variants of each, and all 
repeated over and over. On September 21, he saw a bird give a very 
different, sweet, soft, almost formless, warbled song—a subsong, per- 
haps by an immature individual. A rattling chatter tit-tit-t-t-t-treet, 
sometimes shortened to tit-teerrrt was often heard by Eisenmann (tape 
recorded by E. S. Morton, Cornell Laboratory of Ornithology). 

H. Loftin reports that 1 banded at Cerro Punta March 26, 1967, was 
recaptured March 27, and April 28, 1968, and March 1, 1970. 


HENICORHINA LEUCOPHRYS LEUCOPHRYS (Tschudi ) 


Troglodytes leucophrys Tschudi, Arch. Naturg., 10 Jahrgang, bd. 1, 1844, p. 282. 
( Peru.) 


Characters.—Throat dull grayish white, very faintly, or not at all, 
lined with darker gray; crown slightly more brownish black; under- 


surface somewhat lighter grayish white. 
A female, taken on Cerro Mali, Darién, February 24, 1964, had the 


FAMILY TROGLODYTIDAE 117 


iris reddish brown; basal area of mandible dark neutral gray; rest of 
bill black; tarsus and toes fuscous-black; claws fuscous-brown. 

Measurements.——Males (10 from Cerro Azul and the Tacarcuna 
massif, Panama, and western Antioquia, Colombia), wing 54.4-60.0 
(56.6), tail 26.7-30.5 (28.8), culmen from base 15.7-19.6 (17.9), tar- 
sus 23.7-25.1 (24.8) mm. 

Females (10 from Darién, and from Antioquia and Cauca, western 
Colombia), wing 50.8-55.1 (52.5), tail 25.1-27.8 (25.8), culmen from 
base 15.8-17.4 (16.7), tarsus 22.0-23.8 (22.8) mm. 

Resident. Found locally on Cerro Azul, eastern Province of Pan- 
ama; fairly common on Cerro Tacarcuna and on Cerro Pirre, Darién. 
Elsewhere, it occurs on the slopes of the Andes of Colombia, Ecuador, 
and Peru. 

The Gray-breasted Wood-Wren, less abundant than the white- 
breasted species, is found in mountain forests, possibly more commonly 
than the few records available indicate. On Cerro Azul (March 29, 
1955) I found 1 specimen at 850 m elevation, in the more humid forest 
on the high slopes where the damp winds from the north and east swept 
across the summit. One had been taken here earlier on March 25, 1911, 
by E. A. Goldman. The bird may be more common there than these re- 
ports indicate. Ridgely (1m litt.) notes that he and others have found it 
on Cerro Azul in recent years. 

In the extensive forests on Cerro Tacarcuna, Darien, this species was 
fairly common. In February and March 1964, I collected 1 at the lower 
level camp, and others higher up, near the summit, at 1400 m. On Cerro 
Pirre, Goldman, from March to early June, found them more common, 
as he prepared 8 for specimens, taken at 5000 m near the head of Rio 
Limon. 

The population found in the mountains of eastern Panama is identi- 
fied as the nominate subspecies lJeucophrys, with its type locality in 
Peru, in accordance with the detailed studies of Hellmayr (Journ. fur 
Ornith., 1903, p. 531, and Cat. Birds Amer., pt. 7, 1937, pp. 262-264). 


CYPHORHINUS ARADUS (Hermann); Song Wren, Cucarachero Cantor 
Myrm|[ornis|] arada Herman, Tab. Aff. Anim, 1783, p. 211 (note r). 


Medium size; bill strong, compressed from side to side, with slightly 
raised base above nostrils; throat, upper breast, and side of head chest- 
nut; upper surface very dark brown; wings and tail heavily barred 
with black. 

Description.—Length 115-130 mm. Upper surface deep brown to 


118 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


blackish brown; wings and tail heavily barred with black; sides, fore- 
neck, and upper breast chestnut; rest of undersurface dull grayish 
brown; undertail coverts in one race chestnut. Rarely, an aberrant in- 
dividual may have the foreneck and upper breast partly or entirely pure 
white. 

These widely distributed wrens average slightly larger in body than 
other forest wrens, and also have heavier bills. Perhaps on account of 
this slightly stronger form, they seem somewhat more aggressive than 
related species, and so are seen more easily. They are also more ter- 
restrial than any other wren besides Microcerculus. 

The species ranges chiefly in the humid lowlands, along the Carib- 
bean slopes of southern Honduras, Nicaragua, Costa Rica, and Panama, 
reaching the Pacific slope in humid hill country of northwestern Costa 
Rica and of western Province of Panama (Cerro Campana) and locally 
extending over the Pacific slope in the lowlands of eastern Panama. It 
is also found in Colombia on both slopes and widely in Amazonia. 
Populations from Panama northward, western Colombia, and the Pa- 
cific slope of Ecuador are often separated, as C. phaeocephalus, from 
the C. avadus group found east of the Andes (Hellmayr, Cat. Birds, 
Aine pt, 7, 1934; p, 292). 


CYPHORHINUS ARADUS INFUSCATUS Zimmer 


Cyphorhinus lawrencu infuscatus Zimmer, Amer. Mus. Nov., no. 573, Oct. 11, 
1932) p23. (Carnlloy Costa Rica) 


Characters.—Darker throughout than the race Jawrenciu, found else- 
where throughout Panama; dorsal surface blackish brown throughout, 
with the forecrown in some blacker; chestnut-brown on side of head, 
foreneck and upper breast distinctly darker; rest of undersurface, in- 
cluding the under surface of the wings, darker. 

Measurements.—Males (10 from Bocas del Toro and Costa Rica), 
wing 61.6-65.7 (63.7), tail 29.2-33.2 (30.8), culmen from base 19.4-21.6 
(20.2); tarsus 23.3-29.2 (24.5) mm. 

Females (10 from Bocas del Toro and Costa Rica), wing 56.5-62.2 
(60.6), tail 26.0-29.8 (27.8), culmen from base 18.2-22.0 (19.7), tar- 
sus 22.2-24.2 (23.6) mm. 

Resident. Fairly common in damp woodland in western Bocas del 
Toro, from the valley of the Rio Sixaola to near Almirante (mouth of 
the Western River); and inland above the head of the Rio Changuena 
to 730 m. 

This race has been found especially in the forests bordering Bahia 


FAMILY TROGLODYTIDAE I1g 


Almirante, behind the mangrove swamps, and over the higher slopes of 
the hills. Elsewhere, this race is reported only from Costa Rica. The 
birds range in pairs or in small groups generally near or on the ground, 
and sometimes into the trees. In general habits they resemble the race 
lawrencu, widely distributed through central and eastern Panama. 

On the Caribbean slope of Costa Rica, Skutch found the globular 
nests, with tubular entrances, in woodland in the crotch of a sapling 
from one to several meters above the forest floor. In February one held 
two eggs “which were white, with brown speckles forming a wreath 
about the thick end.” This species also builds dormitory nests in which 
members of a band sleep; they are of the same shape as breeding nests. 

The vocalizations of this population are essentially like those of law- 
rencii, combining guttural phrases and throaty chatter and clucks with 
musical whistles. 

A male taken at Almirante by D. Hicks (Gorgas Memorial Labora- 
tory Collection) October 3, 1964, weighed 24.3 g. 


CYPHORHINUS ARADUS LAWRENCII Lawrence 


Cyphorhinus Lawrencu, “sp. nov. Scl. M.S.,” Lawrence, Ann. Lyc. Nat. Hist., 
New York, vol. 8, May 1863, p. 5. (Atlantic slope, line of Panama Railroad, 
Canal Zone, Panama.) 


Characters.—Paler in color above and on the lower surface; some- 
what lighter brown on crown and back; lighter underneath in chestnut- 
brown of foreneck and upper breast, and lighter, more brownish gray 
on rest of undersurface. 

In an adult male, collected at Juan Mina, Canal Zone, January 12, 
1961, the iris was wood brown; bill wholly black; tarsus, toes, and claws 
fuscous. In another male, at Armila, San Blas, February 21, 1963, the 
iris was dark brown; bill black; tarsus, toes, and claws dark olive- 
brown. 

Measurements.—Males (10 from Colon, Province of Panama, and 
Canal Zone), wing 61.7-66.8 (64.0), tail 27.4-34.1 (30.2), culmen from 
base 19.2-22.2 (20.4), tarsus 23.1-24.8 (24.1) mm. 

Females (10 from Colon, Province of Panama, and Canal Zone), 
wing 58.8-64.1 (61.2), tail 25.3-29.7 (27.1), culmen from base 18.7- 
21.0 (19.8), tarsus 21.0-25.5 (23.4) mm. 

R. S. Crossin recorded the weight of an adult male taken August 9, 
1968, near Gamboa, as 27.4 g. One weighed by Strauch (Bull. Brit. 
Orn. Club, 1977, p. 64) was 24.6 g. 

Resident. Fairly common, on the Caribbean side from the valley of 


120 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


the Rio Indio in northern Coclé and western Colon, through the north- 
ern Canal Zone, base of Cerro Bruja, and eastern Colon, eastward on 
the Caribbean slope to the Colombian boundary, and on the Pacific 
slope in humid hill country from Cerro Campana, western Province of 
Panama, and in eastern Province of Panama from Cerro Azul and 
Chepo, the Rio Bayano Basin to the lower Rio Sambu, Cerro Pirre, and 
the base of Cerro Tacarcuna, Darién In the Canal Zone, while wide- 
spread in canopied woodland of the Chagres Basin and the Caribbean 
coastal area, it is also common nearer the Pacific in the Madden Forest 
Preserve (Ps) ©) Chapelle i isenmann et alo 

These attractive wrens range regularly in pairs and also frequently 
in small groups of half a dozen or more individuals. They are birds of 
the forest, both original and second growth (rastrojo), ranging mainly 
over the ground, but also above it in low undergrowth. Their rapid, 
harsh-toned calls, mixed with whistled notes, as well as their loud songs, 
draw attention. Usually they may appear briefly on low perches, or in 
small open spaces. The groups regularly include families of adults and 
young. This species 1s sometimes seen about army-ant swarms. 

On Barro Colorado Island in March, Skutch (Auk, 1940, p. 300) 
found them sleeping in rounded ball-shaped nests in family groups of 
adults and their grown young. A nest recorded by Jewel, May 7, 1911, 
near Gatun, Canal Zone, was “found in low wet forest two feet from 
the ground .. . consisted of a long tube or tunnel with the nest proper at 
the far end built of sticks, twigs and dead leaves, lined with grasses. 
Eggs, two ... white, very finely and sparingly speckled with brown, 
one almost immaculate.” (Stone, Proc. Acad. Nat. Sci. Philadelphia 
vol. 70, 1918, p. 272.) The eggs measured 16.5 23.3 and! ite 22 16 
mm. Major-General G. R. Meyer found two nests September 1, 1941, 
in the Forest Reserve, Canal Zone. These were bulky and gourd- 
shaped, suspended in the fork of low saplings 125 mm above the ground 
and made of fine grasses lined with the skeletons of dried leaves. One 
contained two eggs that measured 16.5 X 24.6 and 16.7 24.1 mm. Three 
eggs in the other nest ranged from 16.2-16.5X23.2-24.6 mm. J. E. 
Ambrose, on April 6, 1961, found a nest being constructed at Gatun, 
2.3 m above the ground in a sapling. 

The first report of this wren in Panama was by George N. Lawrence 
(Ann. Lyc. Nat. Hist. New York, vol. 7, 1861; p. 293), who recorded 
a specimen received in the first collection made by James McLeannan, 
under the name “Cyphornis cantans (Gm.).” In the reference for the 
account of this wren Lawrence explained that when he found that the 
identification as cantans was in error, he sent the specimen to Mr. 


FAMILY TROGLODYTIDAE I21 


Sclater, “who decided that it was undescribed, and complimented me by 
conferring my name upon it.” 

A common call is a harsh, guttural cutta, cutta, cutta. This vocaliza- 
tion is often intermixed with clear, sweet whistles in its song, which D. 
Harrower syllabized as “percutta wheet, per-cutta whoot, per-cutta 
wheet, per-cutta woot; per-cutta-cuttle wheet whoot; per-cutta whoot 
whoot” (Eisenmann, Smiths. Misc. Coll. 117, no. 5, 1952, p. 47). An- 
other version heard on Barro Colorado Island is ong cutta cutta, whong 
cutta glut, WHOO HEE, etc. (Willis and Eisenmann, Smiths. Cont. 
Zool. no. 291, 1979, p. 25). Still another common vocalization is a 
sweet, rhythmic, slow, whistled, repeated tock, tee, sometimes preceded 
or followed by the nonmusical cutta notes (Eisenmann). F. O. Cha- 
pelle (in litt. to Eisenmann) reported a three-note song without any 
cuttas, “which exactly matches the ‘do’ and ‘fa’ of Guido’s scale’; three 
variations were Do-do-fa; fa-do-Do; and do-Do-do, the capitalized Do 
being an octave higher than the uncapitalized. 

During admittedly short stays in northern Venezuela where Eisen- 
mann saw or heard (vocalizations identified by P. Schwartz) popula- 
tions belonging to the true C. aradus group, he never heard either the 
cutta or the tock, tee vocalizations so characteristic in Panama. 

Morton (in Temple, Endangered Birds, Univ. of Wisconsin Press, 
1977, pp. 379-384) has reintroduced 7 birds of this species to Barro 
Colorado, where it had disappeared between 1960 and 1970. Two of 
these pairs nested successfully within 5 months of their release. 


MICROCERCULUS MARGINATUS LUSCINIA (Salvin): Whistler Wren, 
Cucarachero Ruisenor 


Microcerculus luscima Salvin, Proc. Zool. Soc. London, 1866, p. 69. (Santa Fé, 
Veragua.) 

Microcerculus acentetus Bangs, Proc. New England Zool. Club, vol. 3, Jan. 30, 
1902, p. 56 (1524 m, Boquete, Chiriqui.) 


Small, wrenlike in body form, but with long bill and long legs, and 
unusually short tail on which the tip barely projects beyond the wings. 

Description.—Length 98-110 mm. Upper surface plain deep brown, 
somewhat duller on the crown and hindneck where the feathers are 
edged very narrowly with dusky; wings and tail dull blackish brown, 
the coverts edged narrowly with deep brown, with a small spot of dull 
buff on the tip; side of head brownish gray, becoming grayish white on 
chin and upper throat; upper sides and breast brownish gray; feathers 
of lower breast and abdomen paler brownish gray, with centrally black- 
ish, more or less V-shaped markings, and edged rather widely with dull 


I22 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


grayish white, producing the effect of dark lower underparts with pale 
and variable indistinct scalloped pattern. 

Immature, with pattern of markings of undersurface less distinct. 

Iris varying from dull mouse brown to reddish brown; bill, with 
maxilla and distal third of mandible dull black; gape and lower half of 
mandibular rami dull marguerite yellow in some; in others, bill dark 
neutral gray; tarsus, toes, and claws dull neutral gray. 

Measurements.—Males (10 from Panama), wing 55.8-59.1 (57.7), 
tail 17.9-20.7 (19.5), culmen from base 19.1-21.7 (20.1), tarsus 20.4- 
ZS). 44055.) iaotone 

Females (9 from Panama, with 1 from Costa Rica), wing 51.2-57.0 
(54.1), tail 17.4-21.9 (19.5), culmen 18.0-21.7 (19.8, average of 9), 
tarsus 21.2-23.8 (22.2) mm. 

Resident. Recorded in forested areas throughout the Republic from 
Chiriqui to San Blas and Darieén. 

I. G. Stiles (Wilson Bull. 1983, vol. 95, p. 169) argues persuasively 
that M. philomela, ranging from southern Mexico to central Costa Rica, 
is a species distinct from Juscinia of southern Costa Rica and Panama 
(and the South American populations treated as races of M. margina- 
tus), readily separable by color in adults as well as by voice, as charac- 
terized by Slud. Adults of /uscimia can be distinguished from philomela 
by grayish white throat and more mottled underparts. Stiles suggests 
that until more is known of the relationship of Juscinia to the South 
American populations, and especially to the white-breasted I. margi- 
natus (type from “Bogota,” Colombia, presumably western Amazonia ) 
the population of southern Costa Rica and Panama be tentatively treated 
as a species, M. luscinia, to be called Pale-throated Wren. Eisenmann 
points out that the thin, whistled vocalization of /uscinia is also given 
by populations in western Colombia, northern Venezuela, Ecuador, and, 
including white-breasted populations, in parts of Amazonian Peru, but 
that in southeastern Peru (Madre de Di0s) and adjacent Bolivia, a 
musical song differing from those of both philomela and luscimia is 
uttered (fide B. Coffey, Ridgely, J. Fitzpatrick) by white-breasted 
birds, currently considered to belong to the same subspecies (margina- 
tus), that sing like /uscima farther north. 

As the luscimia song does not remotely suggest that of a nightingale 
(nor does the bird look like that Palearctic bird) Eisenmann thinks 
Whistler Wren might be a more useful name for the southern popu- 
lations, reserving Nightingale Wren for M. philomela. 

These curious wrens are true forest inhabitants found from sea level 
to high in the mountains, where there is suitable cover and reasonable 


FAMILY TROGLODYTIDAE 123 


freedom from disturbance. Blake recorded 1 from 3000 m on the vol- 
cano in Chiriqui. My own few records there were from 1500 m. I 
found them also in the higher forests on Cerro Campana, and on Cerro 
Azul, and to 1300 m on Cerro Tacarcuna. 

This small and secretive species is encountered on the ground in heavy 
forest, moving about under dense cover. On first sight, their rather 
heavy bodies, long bills, and very short tails briefly suggest tiny rails 
rather than wrens. Rarely, in some heavily shaded area, one may ap- 
pear briefly from a tangle of roots, with body bobbing excitedly, utter- 
ing low, high-pitched chattering calls. To Slud and Ridgely its gait 
suggests a Spotted Sandpiper’s. 

Although very infrequently seen, on the basis of its distinctive vocali- 
zation, it proves not uncommon in forest and canopied second-growth 
woods in humid regions, while somewhat local. Between the years 1965 
and 1976 it has been recorded on the basis of voice in the western Chiri- 
qui highlands (between FE] Volcan and Sereno, Pujals, 1976), Veraguas 
(above Santa Fé near and below the Continental Divide (Eisenmann, 
Pujals, N. G. Smith, Ridgely, 1973-75; in 1926 R. Benson collected 1 at 
Santa Fé as low as 480 m [specimen in American Museum of Natural 
History|); Province of Panama (Cerro Campana, Cerro Azul-Cerro 
Jefe hill forest, Rio Bayano Basin forest in lowlands along FE] Llano- 
Carti road, Rio Majé, and “Cuipo” (Cavanillesia) forest on ride along 
road to Jesus Maria (same observers and years); Canal Zone (Chagres 
Basin, near Summit, and Caribbean coastal area); extreme eastern 
Darien (in Tuira basin near Rio Mono to Cerro Quia up to 750 m, 
Ridgely 1975; specimen in Gorgas Memorial Laboratory collection 
from Cerro Quia). Tape recordings have been taken by E. S. Morton 
(Cornell Laboratory of Ornithology) in Panama Province and the 
Canal Zone, and in most areas birds have been mist-netted frequently. 
Specimens have been preserved from Cerro Campana and the Pipeline 
area near Gamboa. 

The distinctive vocalization consists of a series of ten or so long, 
drawn, very thin, high whistles each on one tone, and separated by in- 
creasing intervals of silence, usually from ca. 2-10 seconds, but when 
the song is long perhaps to 15; the last may be separated by as much as 
30 seconds (fide R. Smart). Each note of this series is about a second 
long, and successive whistles seem on the same or very slightly lower 
pitch. Before the thin whistled series starts, there is an introductory 
phrase of about 4 to 7 short notes, usually given so fast that they may 
seem jumbled, moving up and down scale. At Rancho Grande, Vene- 
zuela, Ikisenmann recognized the same thin, whistled song (race squa- 


124 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


mulatus), but the introductory phrase heard was of only three notes. 
P. Schwartz said that when introducing the first of a cycle of songs the 
Opening phrase was more elaborate. 

The nest and eggs appear to be unknown. Slud observed a bird in 
southwestern Costa Rica carrying food on February 17. A female col- 
lected by Strauch (Bull. Brit. Orn. Club, 1977, p. 64) in the Canal 
Zone area in November was in breeding condition and weighed 22.2 g. 


Family MIMIDAE: Thrashers, Mockingbirds 


A family of about 35 species found in the Americas, ranging from 
southern Canada south through the United States, Central America, 
and the West Indies to southern Argentina and Chile. Two species are 
found in Panama, one of them, the Tropical Mockingbird, was intro- 
duced from Colombia. The other, the Gray Catbird, is a winter season 
visitor from the north. 


KEY, fO SPECIES OF MIMIDAE 


1. Undersurface of body white; crown gray like back. 
Tropical Mockingbird, Mimus gilvus tolimensis. p. 124 
Undersurface of body gray; crown black; back gray. 
Gray Catbird, Dumetella carolinensis. p. 127 


MIMUS GILVUS TOLIMENSIS Ridgway: Tropical Mockingbird, 
Sinsonte del Caribe 


Mimus gilvus tolimensis Ridgway, Smiths. Misc. Coll., vol. 47, August 6, 1904, 
p. 113. (Plains of Tolima, Boyaca, Colombia.) 


Rather large, with long, white-tipped tail; gray above, white under- 
neath. 

Description—Length 240-275 mm. Adult (sexes alike), upper sur- 
face of head and body somewhat brownish light gray; crown narrowly 
streaked with dusky; wings and tail blackish slate, with indistinct nar- 
row pale gray edgings, becoming white on tips of greater coverts where 
they form two somewhat indefinite bands; tertials and secondaries 
edged narrowly with white at distal end; tail tipped broadly with white, 
with this continued as a narrow border on the outermost rectrix to the 
base; a dusky streak from the lores to back of the eye, covering the 
free margins of the eyelids; a somewhat indistinct band of white above 
the eye, becoming broader behind; undersurface white, tinged more or 


FAMILY MIMIDAE I25 


less faintly with gray across breast; axillars and underwing coverts 
white, marked indistinctly with dusky. Immatures are more brownish 
above. 

Iris yellow; bill black; tarsus and toes fuscous. 

Measurements.—Males (10 from the range of the race in Colombia), 
wing 116.2-129.5 (121.7), tail 119.7-128.7 (125.4), culmen from base 
25-251, (23.1, average of 9); tarsus 31.9-35.9 (34.7) mm. 

Females (10 from the range of the race in Colombia), wing 112.1- 
mero Ci7.6), tail 110.2-128.5 (116.1), culmen from base 20:2-25.7 
(23.2), tarsus 31.7-35.8 (33.2) mm. 

Introduced. Common in the Canal Zone and adjacent Colon and 
Province of Panama, on both slopes, in cleared, especially residential, 
areas, east at least to Portobello, Colon, and, on the Pacific slope, to 
Canita, near the Bayano Dam site, where Ridgely (1 litt.) found a 
pair on December 29, 1974. 

Introduced from its range in western and central Colombia, its pres- 
ence is due apparently to birds released through customs control, and 
probably in part also by escape from cages. 

The first definite report of this mockingbird in Panama that has come 
to attention is that of Herbert G. Deignan (Auk, 1933, p. 125) who 
wrote that “On July 13, 1932, while standing in front of the Balboa 
station of the Panama Railroad, I heard the song of a mocking bird 
and discovered the bird perched at the top of a flagpole nearby.”” Arbib 
and Loetscher (Auk, 1935, p. 326) listed the mockingbird from the 
Canal Zone in summer, 1934, but were told (in error) that it had been 
introduced from California. Chapman (Auk, 1941, pp. 98-99), in 
January 1938, recorded a pair at the Gorgas Memorial Institute in 
Panama City, and at Balboa, and mentioned that it was reported at 
Pedro Miguel “breeding” near Ancon. In 1938 and 1939, he secured 
specimens, which he found were Mimus giluus of South America, and 
not M. polyglottos of North America, as some had supposed, referring 
them to the Venezuelan melano pterus. 

Barlier, Tollef Monniche had collected a male, August 23, 1937, at 
the Finca Lérida above Boquete, Chiriqui. This, Blake (Fieldiana, 
Zool. vol. 36, no. 5, 1958, p. 548) in his account of the Monniche collec- 
tion, considered to be a locally-escaped cage bird, and not an extension 
of range from the far distant Canal Zone. The spread had begun, how- 
ever, and in Iebruary 1952, I recorded several and collected a male near 
the mouth of the Rio Indio in western Province of Colon. Here also 
they may have become established through release of cage birds. 


126 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


The male from Lérida, while listed by Blake as M. g. melanopterus, 
on examination proves to be the somewhat larger race tolimensis. 

With regard to introduction, an account received through the ento- 
mologist James Zetek, long resident in Panama, relates that during 
World War Ia shipment of mockingbirds that originated in Medellin, 
Colombia, came to the Canal Zone en route to the United States. Since 
all birds were temporarily prohibited entry into that country as a pre- 
caution against the disease psittacosis (then little understood), the car- 
rier released the entire shipment. 

I was told of a similar occurrence in 1956 by Mr. E. J. Husted, re- 
tired employee under the Canal Zone government and a resident of the 
Canal Zone since boyhood. Mr. Husted related that in late 1935, about 
100 live mockingbirds in cages were brought from Colombia on a ship 
that docked at Pier 18 in Balboa. They were loaded on a truck in Mr. 
Husted’s charge, the birds being intended for sale in the market. When 
the authorities demanded a dollar import duty per bird, the importer, 
who had expected to sell them at $1.50 each, became disgusted and re- 
leased the entire shipment. 

Colombia has 2 populations of this mockingbird that differ slightly 
in size. The somewhat larger form, the subspecies Mimus gilvus toli- 
mensis, found in western and southern Colombia, measures, as follows: 
Male, wing 116.2-129.5, tail 114.7-128.7 mm. Female, wing 108.9- 
122.3, tail 102.4-120.4 mm. The other, Mimus gilvus melanopterus, 
slightly smaller, has the following dimensions: Male, wing 109.1-119.8, 
tail 102.4-120.4 mm. Female, wing 93.4-101.2, tail 100.1-108.3 mm. 
It is found in northeastern and eastern Colombia to Venezuela and 
Guyana southward to adjacent northern Brazil. 

The two are closely similar in pattern of markings and in color. 
While it is possible that both may have been represented in imported 
captives, the few that have been available for examination as museum 
specimens have the size of the larger tolimensis. 

In Ancon, where I have seen them regularly, they carry nesting ma- 
terial into the tops of the royal palms in late December and early Janu- 
ary. Young with short tails, recently from the nest, have been observed 
from mid-December to mid-September. 

In song and mannerisms this mockingbird is closely similar to our 
common species Mimus polyglottos of the United States, although it is 
not known to imitate other birds. On the ground, males frequently 
stand or move with partly spread, waving wings (which are plain, with- 
out white), exactly like our northern species. The mannerism is one 


FAMILY MIMIDAE 127 


that I have found especially interesting in view of the arguments as to 
why our northern bird with the prominent white patch on the wing has 
this habit. It is evident that the display is in the wing action, with no 
basis in color pattern. 


DUMETELLA CAROLINENSIS (Linnaeus): Gray Catbird, Pajaro Gato 
Muscicapa carolinensis Linnaeus, Syst. Nat., ed. 12, vol. 7, 1766, p. 328. ( Virginia.) 


Medium size, slender, with long tail; dark gray. 

Description.—Length 185-215 mm. Adult (sexes alike), crown, 
wings, and tail black; wings, and to a lesser degree, tail, margined with 
gray; body slate gray, darker above, paler below; undertail coverts 
chestnut. 

Bill black; tarsus and toes blackish brown; iris dark brown. 

Measurements.—Males (10 from the breeding range), wing 86.0- 
96.0 (91.2), tail 89.0-103.0 (95.9), culmen from base 20.0-20.2 (20.1), 
tarsus 27.0-29.0 (27.9) mm. 

Females (10 from the breeding range), wing 84.0-91.0 (88.1), tail 
82.0-97.0 (91.7), culmen from base 19.2-20.4 (19.8), tarsus 27.0-28.5 
(27.9') mam. 

Migrant from temperate North America; common in Bocas del Toro, 
September 1-March 31 (May 29, once), (Almirante, Changuinola, 
Cocoplum). Occurs casually and irregularly elsewhere in western Pan- 
ama, in Chiriqui ( Boquete), and Veraguas ( Paracoté); winters in very 
small numbers and irregularly to the Canal Zone chiefly on the Carib- 
bean slope (Curundu, Corozal, Coco Solo, Barro Colorado Island, Juan 
Mina, Frijoles, Ft. Davis, Gamboa Pipeline Road, Gatun, Lion Hill); 
casual in San Blas (Mandinga, Armila, Puerto Obaldia); recorded on 
Islas Escudo de Veraguas and Bastimentos, and on Taboga Island in 
the Gulf of Panama (Sturgis). On the Pacific slope the farthest east 
appears to be one seen by Ridgely at Cerro Azul on March 11, 1979. 

The catbird is common in thickets and low forest around Bahia Almi- 
rante, ranging usually under cover, calling only occasionally. Though 
not especially timid, they usually remain under leafy cover so that rela- 
tively few are seen. Rarely, one may sing. Their true number becomes 
evident when mist nets are set, as many are captured. 

Dr. Pedro Galindo, who has netted catbirds regularly in his studies 
at Almirante, has furnished the following interesting information. One 
that he banded October 18, 1963, was reported from Oxford, Nova 
Scotia, in July 1964; and another, caught December 19, 1963, was 


128 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


captured near Springfield, also in Nova Scotia, July 4, 1964. Four 
others given bands near Almirante were recaptured subsequently at 
that banding locality as follows: one November 6, 1963, was retaken 
there October 13, 1967; one on October 13, 1964, was retaken October 
4, 1967; one April 8, 1965, was recaptured April 13, 1967; and one on 
January 16, 1964, was recaptured July 11, 1967. The returns from 
Nova Scotia mark the far northeastern border of the breeding range of 
the species, while Panama and Colombia are the southern limit of the 
wintering range. 

In another netting study, Rogers and Odum (Wilson Bul., 1966, p. 
418) found that in Panama newly-arrived southbound migrant catbirds 
that had just flown across the Gulf of Mexico weighed 26.5-37.4 
(31.2) g and had exhausted their fat reserves; some had also experi- 
enced weight loss in the breast muscles, which seemed reduced so that 
the keels of the sternums protruded noticeably. 

Reports of netting around Almirante indicate that many birds winter 
in the area and a number return the following or subsequent seasons. 
The migration generally begins in early October and the peak is reached 
the fourth week of that month, with a few birds remaining into May 
(Galindo and Méndez, Bird-Banding, 1965, pp. 233-234; Loftin, Carib. 
Journ. Sci., 1963, p. 66; Loftin and Olson, Carib. Journ. Sci., 1964, p. 
194). 


Family TURDIDAE: Thrushes, Zorzales 


The thrushes are a worldwide family of approximately 300 species 
that include some of the most familiar birds of temperate regions, 
known for their confiding presence around human habitation and for 
their fine vocal abilities. Fourteen species are found in Panama, in- 
cluding 4 migrants from the north. The native species fall into two 
groups, robins of the genus Turdus, and nightingale-thrushes, genus 
Catharus. The robins have habits similar to the North American Tur- 
dus migratorius, but those inhabiting dense forest are sometimes hard 
to see—although they will often approach a squeaking sound. The 
Clay-colored Robin is widespread in the lowlands and at home in yards 
and gardens; the other species of Turdus are found from the foothills 
to above timberline. Robins usually place their cup-shaped nest in a 
tree or bush. The Clay-colored Robin, like its northern relative, has so 
accustomed itself to human habitation that it sometimes builds its nest 
in the angle of a roof. 


FAMILY TURDIDAE I29 


The clear flutelike phrases of some of Panama’s nightingale-thrushes 
are among the most beautiful songs produced by any bird. The night- 
ingale-thrushes are highland birds, found mainly in the west. They 
nest on or near the ground. Like the robins, their diet usually is mostly 
fruit or animal matter. 

The solitaires, M/yadestes, have long been considered part of the 
thrush family (Ripley, Check-list Birds World, 1964, p. 89), but recent 
evidence assembled by Sibley (Auk, 1973, p. 408) and Ames (Bonn. 
Zool. Beitr., vol. 26, 1975, p. 127) suggests that they are more closely 
‘related to the silky flycatchers, Ptilogonatidae. I have followed their 
recommendations in including Myadestes in the Ptilogonatidae and 
placing that family next to the Turdidae. 


KEY TO SPECIES OF TURDIDAE 


SoS Me LESS) UleAT AL OC) ata Liao idee Glace a eee ee ate itke ci Somes ewe Payons 2 
Penccmenione «liaiay lOO’ mag oy 3. Aton ios ae oe Res Sheba een es 10 
PMO IP OGatT@aACETS POLLO eth iA oc eos oldu alee WME are eun engl tale 5 3 
AU MOE SUIGhACE UIMSDOLLEU ke semi 4). crs le ccs take Abe as Guedes ened Gh clwecne 6 


3. Spotting extensive, from throat to abdomen. 
Wood Thrush, Hylocichla mustelina. p. 143 
Spocme restricted to’ throat and upper chest... 2.02. Sere oe. ene 4 
4. Eye-ring distinct. 
Swainson’s Thrush, Catharus ustulatus. p. 144 
Fieve gl MMMM ISGIIl CLYOT tANSEMt eyes ost ses cies aca cue ae ac ee et okle : 5 
5. Spotted area of upper chest washed with cinnamon-buff. 
Veery, Catharus fuscescens. p. 149 
Spotted area of upper chest whitish. 
Gray-cheeked Thrush, Catharus minimus minimus. p. 148 
6. Back blackish-gray. 
Slaty-backed Nightingale-Thrush, Catharus fuscater. p. 154 


IEXSAGUS IDNRONR TDS gael Wk 8 es in la Aa uta aS i a a a cole Te be I ed 7 
Sera mmebrowilor DiaCkK. sa hia eee eo le Pees ee bee ee. 8 
CAROTID» GIP 53 hoe sini Nin LIBi so ZC Ue lies 8 a sr eC US ee ae 9 


8. Crown brown. 
Ruddy-capped Nightingale-Thrush, Catharus frantzi wetmoret. p. 158 
Crown black. 
Black-headed Nightingale-Thrush, Catharus mexicanus fumosus. p. 152 
9. Chest with greenish-brown band. 
Black-billed Nightingale-Thrush, Catharus gracilirostris. p. 163 
Chest gray. 
Orange-billed Nightingale-Thrush. Catharus auranturostris. p. 160 
10. Throat boldly striped dusky and white, upper chest white. 
White-throated Robin, Turdus albicollis. p. 130 
Mineat striped indistinctly or Mok at alls. eee Geese ee woe ee heck ce 11 


130 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


1. Undersuriace: brownie uhh Eee Ce 12 
Undersurface exay on blackish i ea 13 
12. Upper surface dark brown. 
Pale-vented Robin, Turdus fumigatus obsoletus. p. 135 
Upper surface light brown. 
Clay-colored Robin, Turdus grayi casius. p. 137 


13. Undersurface gray. 
Mountain Robin, Turdus plebejus plebejus. p. 140 
Undersurface blackish. 
Sooty Robin, Turdus nigrescens. p. 142 


TURDUS ALBICOLLIS Vieillot: White-throated Robin, 
Casca Gargantiblanco 


Turdus albicollis Vieillot, 1818, Nouv. Dict. Hist. Nat., nouv. ed. 20, p. 227. 
(Brazil = Rio de Janiero.) 


Rather large; robinlike; upper throat boldly streaked dusky and 
white, upper chest pure white. 

Description.—Length 190-226 mm. Adult (sexes alike), upper sur- 
face from crown to rump brownish olive; scapulars, wing coverts, and 
secondaries brownish olive; primaries and rectrices darker brown; side 
of face brownish olive; throat boldly streaked dusky and white, with 
white continuing on to upper chest; rest of undersurface tawny-olive, 
to abdomen and undertail coverts, which are white or pale buff; under- 
wing coverts tawny-olive. 

Juvenile, scapulars and feathers on upper back veined and tipped 
russet; undersurface from throat to breast, and sometimes to abdo- 
men, spotted dusky. 

The White-throated Robin is a wide-ranging species, found from 
northern Mexico through Central America and into South America as 
far as southernmost Brazil, Paraguay, and northern Argentina in Mis- 
iones Province. In Panama it occurs mainly on the Pacific slope, in 
Chiriqui, Veraguas, the western side of Azuero Peninsula, in Coclé at 
El Valle, and eastern Province of Panama at Cerro Campana, and in 
eastern Darién. There are several recent sightings from Barro Colo- 
rado Island after the nesting season (Ridgely, 1976, p. 274) and else- 
where in the Canal Zone (Toucan, May 1977, pp. 3-4). This species 
1s common in forest and forest borders in the foothills and lower high- 
lands. Aldrich and Bole (Scient. Pub. Cleveland Mus. Nat. Hist., vol. 
7, 1937, pp. 25, 26) found it common in rain forest and cloud forest 
(900 m and above) on the western slope of the Azuero Peninsula, and 
Worth (Auk, 1939, pp. 307-308) called it common at 168 m in forests 
and clearing near Rio Gariché, Chiriqui. The population on Coiba Is- 


FAMILY TURDIDAE P31 


land, subspecifically distinct from the mainland forms, lives at sea level. 

All observers have commented on the similarity of the usual song of 
this species and T. migratorius. The call, however, is entirely different 
—a rather froglike guttural, harsh wmk or yuck. It also gives a screechy 
kyeé-ee-yoo somewhat suggestive of the whining call of T. gray. It 
also, more rarely, gives a rich, repetitive song more suggestive of a 
mimid or wren than a robin. 

Sometimes seen singly or in pairs, I have also found White-throated 
Robins feeding in fruiting trees in groups of a dozen or more. Its diet 
is mainly vegetable. Some of E. A. Goldman’s notes of stomach con- 
tents of specimens of T. a. daguae collected at Cana, Darién, are typi- 
cal: one with the stomach two-thirds full had bits of a spider 3%, 5 
drupes of Ficus sp. 85%, wild fruit skin 12%; one a fourth full had 
the heads of two carabid larvae 5%, 2 seeds of an unidentified plant 
and other fragments 65%, 1 drupe of Oleaceae sp. 30%; and one with 
the stomach nearly empty had bits of a locustid 15% and bits of skin 
of a wild fruit 85%. Willis (Living Bird, 1966, p. 202) has found 
single individuals following army-ant swarms on Barro Colorado Is- 
land September through January. 


TURDUS ALBICOLLIS CNEPHOSUS (Bangs) 
Merula leucauchen cnephosa Bangs, 1902, Proc. New England Zool. Club, 3, p. 92. 

(Boquete, Volcan de Chiriqui, Panama.) 

Characters.—Upper surface brownish olive; bill yellowish. 

A female taken March 5, 1951, at Cerro Campana, Panama had the 
iris light brown; prominent thickened eye-ring honey yellow; base of 
maxilla and tip of culmen dull dark brown; rest of bill light greenish 
olive; tarsus dull greenish olive; toes light brownish white. 

Measurements.—Males (10 from Chiriqui and Coclé), wing 114.6- 
ae V9.5), tail 85.2-96.2 (90.3); culmen from base 21.3-23.7 
(22.7), tarsus 28.6-31.1 (30.0) mm. 

Females (10 from Chiriqui, Coclé, and Province of Panama), wing 
114.7-124.8 (119.1), tail 82.4-93.2 (88.7), culmen from base 22.0- 
24.0 (22.9, average of 9), tarsus 29.8-32.0 (30.7) mm. 

Resident. Fairly common in Chiriqui, Veraguas, Coclé to western 
Province of Panama on the Pacific slope in forest and forest edges of 
middle elevations, at least occasionally wandering to lowlands. It is 
also found in southwestern Costa Rica. Aldrich and Bole (Scient. 
Publ. Cleveland Mus. Nat. Hist., vol. 7, 1937, p. 114), on the western 
slope cf the Azuero Peninsula, said “this species was first encountered 
at Mariato River camp at 250 feet elevation, but became more abundant 


132 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


as we ascended in the mountains. It reached its peak of abundance at 
our highest camp, Cavulla, near the top of Cerro Viejo at 3000 feet 
elevation,” in subtropical forest. 

I have encountered this thrush in many localities in Chiriqui: on 
February 26, 1954, I shot 2 males at El Volcan in open forest at the 
edge of the llano when the woods were completely dry; on March 8, 
1954, I noted several in forest at Palo Santo; on March 27, 1954, I took 
1 at Laguna Grande; the next day I found others singing and calling in 
the tops of tall trees at 1490 m in Silla de Cerro Pando; several were 
in heavy woodland on the Quebrada Candela at Sereno on February 
23, 1955; at San Félix I shot 1 on February 19, 1956; and at Boquete I 
found several in forest at 1100 m. 

In Coclé I have collected specimens at El Valle from Rio Anton, 
from Cerro La India Dormida at 750 m, and from the open forest along 
the head of Rio Mata Ahogada (480 m), where it was common. 

Arcé collected it in Veraguas at Santa Fé (not dated). 

In the Province of Panama I found a dozen or so on Cerro Campana 
in a tract of heavy forest where a purple berry was common. They re- 
mained under heavy cover and responded readily to a squeak. A male 
collected by Strauch (Bull. Brit, Orn. Club; 1977, p: 64)ar Geum 
Campana weighed 66.2 g, a female 77.2 g. 

The song of the White-throated Robin is usually described as robin- 
like and very fine, consisting of rich, powerful phrases mixed with 
shorter phrases that are high and weak. In Costa Rica, Skutch (Pac. 
Coast Avif., no. 34, 1960, p. 84-85) has heard them singing from early 
February until August 20, with a peak of song in early April. There is 
also another non-robinlike song less commonly heard. Its calls include 
a peculiar note like the high-pitched croak of a frog, which does not at 
all suggest a Turdus. 

In Panama, nests and eggs are known from April through July. 
Blake (Condor, 1956, p. 387) describes a nest collected April 28, 1932, 
by Monniche at approximately 1620 m on the slopes of Volcan de 
Chiriqui that was located half a meter above the ground in a large rot- 
ten stump. The nest was a shallow cup 76.2 mm wide and 38.1 mm 
deep, loosely constructed of coarse twigs. It contained 3 partly incu- 
bated eggs “‘so heavily speckled and blotched with dull reddish-brown 
as largely to obscure the pale greenish-white ground cover.” The 
eggs measured 31.3 20.6, 30.4 20.6, 31X21 mm. 

Worth (Auk, 1939, p. 307) found a nest along the Rio Gariché, 
Chiriqui (1200 m) that on June 26, 1937, contained 2 eggs with a 
ground color “almost white, tinged with a very pale and diluted green. 


FAMILY TURDIDAE 133 


They were heavily spotted with small dots of russet, so thickly clus- 
tered at the blunt end as to become confluent.” Their measurements 
were 29X21 and 28X22 mm. The nest was nearly 2 m from the ground 
in the crotch of a small coffee tree, with a foundation of coarse dead 
grass and a superstructure of living green moss. The cup-shaped cavity 
was lined with fine rootlets. The eggs from this nest hatched on July 3. 
At hatching the young had a few tufts of long buffy down, and by July 
17 one of the fledglings had already flown from the nest. 

E. S. Morton (im htt. to Eisenmann) reports the White-throated 
~Robin a common breeder on Cerro Campana, western Province of Pan- 
ama, in April-July in 1970 and 1971. Some nests were in niches formed 
by epiphytes or on depressions on the trunk of a tree, and were com- 
posed mainly of moss with a little mud. During most of the dry season 
the species almost completely disappeared, probably descending to the 
humid lowlands or dispersing widely in search of fruit. 


TURDUS ALBICOLLIS COIBENSIS Eisenmann 


Turdus assimilis coibensis Eisenmann, 1950, Auk, 67, p. 366. (Coiba Island, 
Veraguas, Panama.) 


Characters.—Above and below ruddier than cnephosus; undertail 
coverts with broad fuscous margins; bill largely dark. 

A male taken March 23, 1962, on Isla Brincanco, Islas Contreras, 
had the iris orange-brown; eye-ring honey yellow; base of bill dull 
fuscous-black; tip dull greenish yellow shading to honey yellow on 
distal half of cutting edge; tarsus and toes light buffy brown; claws 
dull brownish white. 

Measurements.—Males (8 from Isla Coiba and 2 from Isla Brin- 
canco), wing 109.5-117.5 (112.8), tail 74.8-89.1 (83.1), culmen from 
base 20.5-22.9 (21.7), tarsus 27.6-31.8 (30.1) mm. 

Females (5 from Isla Coiba and 1 from Isla Rancheria), wing 108.0- 
113.4 (112.1), tail 79.5-86.8 (82.6), culmen from base 21.0-23.3 (22.2), 
tarsus 29.0-31.0 (30.2) mm. 

Resident. This is the only thrush on Coiba Island and adjacent Isla 
Rancheria. In January and February 1956 it was the most abundant 
bird on Coiba in high forest, although unlike the mainland forms, here 
it also descends to sea level, where I found it in the edge of a mangrove 
swamp at the Rio Catival. In the Punta Damas section of the island I 
saw them feeding on berries in trees and searching among the dry 
leaves on the trail. By squeaking I often drew ina dozen. Their call is 
a complaining chur-r-r or pru-rr-r, and their song, which I heard on 


134 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


January 22 and 31, suggested T. grayi casius but with a slower delivery 
and notes much higher on the scale. In early April 1976 Ridgely found 
it common in forest, singing all day, a song of excellent quality; it ut- 
tered a guttural call like birds of the western highlands. 

On March 23, 1962, I collected 2 male White-throated Thrushes on 
Isla Brincanco, Islas Contreras. Both birds were in full breeding con- 
dition and several times I heard their sweet robin song. I have tenta- 
tively assigned them to the race coibensis, although with a larger series 
subspecific differences may become clear. Eisenmann notes that bill 
colors of these birds suggest intermediacy between the dark-billed birds 
of Coiba and the yellow-billed western mainland birds. The Coiba race 
of the Streaked Saltator, Saltator albicollis scotinus, is also known to 
extend to Isla Brincanco. 


TURDUS ALBICOLLIS DAGUAE Berlepsch 


Turdus daguae Berlepsch, 1897, Orn. Monatsb. 5, p. 176. (San José, Rio Dagua, 
Colombia. ) 


Characters.—Undersur face sepia; upper surface ruddier than other 
Panama races. 

A male taken January 31, 1961, on Cerro Pirre, Darien, had then. 
bright brown; thickened eyelid dull lemon yellow; maxilla and tip of 
mandible black; base of mandible light olive-green; tarsus and toes 
brownish gray. 

A male taken March 9, 1964, at Tacarcuna Village, Darién, had the 
iris warm brown; thickened edge of eyelid bright, somewhat greenish 
yellow; maxilla fuscous-black; cutting edge of mandible and gape dull 
yellow; end of mandible and extreme base of rami fuscous, middle 
section dull yellowish green; tarsus and toes dull, somewhat grayish 
brown; claws somewhat darker. 

Measurements.—Males (10 from Darién, Panama, and Choco, Co- 
lombia), wing 102.4-110.9 (108.2), tail 73.3-81.6 (77.3, average of 9), 
culmen from base 19.0-21.6 (20.1, average of 9), tarsus 26.8-30.1 
(28.1) mm. 

Females (7 from Darién, Panama, and Choco, Colombia), wing 
103.1-107.4 (105.6), tail 73.0-77.9 (75.4), culmen from base 20.4-21.8 
(21.2), tarsus 27.1-29.6 (27.9) mm. 

Resident. Fairly common in eastern Darién on the Pacific slope. 
Found also in western Colombia and Ecuador. The Smithsonian has 
several specimens taken by E. A. Goldman between February and June 
1912 at Cana, at elevations from 540 to 600 m, and I have collected 


FAMILY TURDIDAE 135 


others at Tacarcuna Village (570 m) and on Cerro Pirre at the head of 
the Rio Seteganti (450 m). The 3 birds I took at Cerro Pirre all had 
the tail feathers damaged or cut, as though they had been feeding in 
“cana brava” (Bactris minor), a small palm heavily armed with long 
slender spines. The male | collected at Tacarcuna Village on March 9, 
1962, was in breeding condition; its song was clear and sweet with 
robin tones, but with many notes that suggested an oriole. 

The only other indications of the breeding season of this race in 
Panama comes from Bangs and Barbour (Bull. Mus. Comp. Zool., vol. 
65, 1922, p. 221), who reported 3 adults and 2 “spotted young” taken 
at the headwaters of the Rio San Antonio on Mt. Sapo in April 1922. 
A heavily spotted immature in the Smithsonian collection (USNM no. 
427026) was collected by Carriker in Colombia at Villa Arteaga, Antio- 
quia, on May 6, 1950. 


TURDUS FUMIGATUS OBSOLETUS Lawrence: Pale-vented Robin, 
Casca Barriga Blanca 


Turdus obsoletus Lawrence, Ann. Lyc. Nat. Hist.. New York, vol. 7, February 
1862, p. 470. (Panama. ) 


Bill black; plain dull brown above; breast and sides somewhat paler. 

Description.—Length 215-225 mm. Adult (sexes alike), upper sur- 
face, wings, and tail warm sepia-brown; breast and sides wood-brown; 
throat brownish white streaked with dull brown; abdomen and under- 
tail coverts white; underwing coverts ochraceous-buff. 

Immature, upper surface somewhat duller; more olive above, with 
pale narrow shaft lines; wing coverts spotted with ochraceous-buff; 
throat somewhat buff-gray, spotted and lined with olive-brown; breast 
and sides dull buff, spotted heavily with blackish brown; abdomen and 
undertail coverts white; underwing coverts as in adult. 

An adult male, taken February 25, 1964, at 1280 m on Cerro Mali, 
Darién, had the iris dark mouse brown; bill black, except for the 
fuscous-colored base of the mandibular rami; tarsus and toes olive- 
brown; claws fuscous. 

Measurements.—Males (10 from Cerro Campana, and Darién), 
wing 119.2-124.5 (121.6), tail 84.3-92.3 (89.3), culmen from base 
23.0-25.3 (24.4), tarsus 28.7-31.0 (30.0) mm. 

Females (10 from Bocas del Toro, eastern Province of Panama, San 
Blas, and Darien), wing 110.4-122.0 (115.9), tail 79.5-90.2 (84.2), 
culmen from base 21.0-25.1 (23.6), tarsus 27.9-31.5 (29.9) mm. 

Resident. Found in forested mountain areas in Bocas del Toro 
(upper Rio Changuena), Chiriqui (reported from the Caribbean slope 


136 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


of Volcan de Chiriqui, which might be the Bocas del Toro side of the 
Continental Divide), Veraguas (Santa Fé), Province of Panama (Cer- 
ro Campana and Cerro Azul), Darién, and San Blas. 

The race obsoletus 1s found also on the Caribbean slopes of Costa 
Rica and in northwestern Colombia (Jiménez). Eisenmann writes: “I 
don’t think the T. obsoletus complex (from Costa Rica to western 
Ecuador) is conspecific with the T. fumigatus of eastern Amazonia 
and northwestern South America—which was Hellmayr’s (Cat. Birds, 
Am., vol. 7, 1934, p. 389) very broad treatment. Gyldenstolpe (Kungl. 
Sv. Vet. Akad. Handlingar, 22, no. 3, 1945, p. 279) concluded that the 
T. hauxwelli complex must be removed from the species T. fumigatus, 
as they proved to be sympatric in Brazilian Amazonia. Ripley (Check- 
list Birds World, vol. 10, 1964, p. 219) followed Gyldenstolpe in the 
specific separation of the T. hauxwelli complex from T. fumigatus. 
Probably through an oversight, he left the T. obsoletus complex in 
IT’. fumigatus when its morphological affinities and geography are with 
the more western T. hauxwelli. Meyer de Schauensee (Birds of South 
America, 1966, pp. 416-417) accordingly merges the T. obsoletus and T. 
hauxwelli complexes in one species, using, because of priority, T. ob- 
soletus as the species name.” In limited encounters I have seen these 
thrushes singly, for brief periods only, in trees or occasionally on the 
ground, usually near or under cover. Most of the specimens that I 
have handled were captured 1n mist nets. This species is a bird of the 
forest interior, and almost entirely arboreal. 

Ridgely (1976, p. 275) describes the song as similar to the Clay- 
colored Robin’s, but faster and less rich. 

The type specimen, without definite locality, has been attributed al- 
most surely erroneously to Lion Hill or the Panama Railroad in the 
Canal Zone (where there is no other record of this species). 

Stomachs that I have examined have been filled with fragments of 
drupes and small seeds, with an ant in one (perhaps swallowed with a 
berry). 

The statement by Ridgway (U.S. Nat. Mus. Bull. 50, pt. 4, 1907, p. 
92) that fumigatus (sensu stricto) of northern South America has 
“much greater adhesion of the anterior toes” than other thrushes does 
not hold in the series examined. 

There has been no report of the nest or eggs of the Panama race. 
Belcher and Smooker (Ibis, 1937, pp. 511-512) described the nest and 
eggs from Trinidad as “a bulky cup with exterior of green moss and 
lining of rootlets. In cacao plantations it is placed about 15 feet up as a 
rule; in forest the site may be in the crown of a tree-fern or near the 


FAMILY TURDIDAE L377, 


top of a slender sapling, or, more rarely, in a recess in the stem of some 
great tree. On roadsides they may be built in a niche or on a projection 
of a steep face, and here they are often higher off the road surface than 
is the nest of T. albicols. The eggs are usually three, less often two; 
they are similar to, but rather larger than, the eggs of T. albicollis; the 
deep blue type with pronounced markings does not, however, occur in 
this species. Typical eggs may be described as regular ovals, smooth- 
shelled, and slightly glossy. The ground is pale greenish-blue, and the 
markings are profuse marblings and blotches of pale reddish-brown, 
converging toward the larger end and there forming a cap. Average 
measurements are 28.2 20.4 mm.: a large set of two measure 32.2X 
20.8 and 3221.4 mm.” 


TURDUS GRAYI CASIUS (Bonaparte): Clay-colored Robin, 
Casca Pardo 


Planesticus casius Bonaparte, Compt. Rend. Acad. Sci., vol. 41, no. 17, 1855 (not 
earlier than October 22), p. 657. (Panama.) 

Rather dull brown in color throughout; a typical “robin” (in the 
American sense) in appearance and action. 

Description.—Length 205-235 mm. Adult (sexes alike), dull clay 
brown to buffy brown on undersurface, with the throat paler and 
streaked with dark brown; darker brown above. 

Juvenile, brighter buffy brown on undersurface, with broken bars of 
dull grayish brown; back and sides of head streaked, and wing coverts 
tipped with cinnamon-buff. 

Iris verona to orange-brown; anterior half of bill, and the cutting 
edge of the mandible, dull honey yellow; basal half dull greenish gray; 
tarsus, toes, and claws dull neutral gray, the claws sometimes buff at 
tip. The bill color is brighter in breeding season. 

Measurements—Males (14 from Panama), wing 114.1-129.3 
(120.3), tail 89.1-106.8 (95.5) culmen from base 22.3-24.4 (23.1), 
tarsus 30.0-33.0 (31.2) mm. 

Females (12 from Panama), wing 110.2-119.2 (115.2), tail 85.9- 
Yeo (922). culmen: from; base 21.1-25.0.(22.9); tarsus 27.9-32.2 
(30.3) mm. 

Resident. Common, chiefly in the Tropical Zone. On the Pacific 
slope from Costa Rica east, including the Azuero Peninsula through 
the Province of Panama (where recorded east to the lower course of 
the Rio Majé). On the Caribbean side, from Bocas del Toro through 
northern Veraguas, Colon, and the Comarca de San Blas to Colombia 
(Acandi, Choco). In Chiriqui it ascends to the Subtropical Zone above 


138 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


KE] Volcan and Cerro Punta, and in Boquete (Lérida, Quiel); recorded 
also near Santa Fé in Veraguas, (elevation not stated), Isla Canal de 
Afuera (sight record); Isla Cébaco. Not recorded from Perlas Is- 
lands nor observed on Isla Coiba by any recent students (a specimen in 
the American Museum of Natural History collected by Batty is prob- 
ably mislabeled, as are many of his “island” specimens). 

The Clay-colored Robin is the best known of its group in Panama 
because of its wide distribution. Under primitive conditions it is com- 
mon in open forest growth, and in brushy areas; in denser stands found 
mainly along streams or other borders. As the land 1s cleared for culti- 
vation this is one of the birds that is able to adapt to the changed con- 
ditions, perhaps even to profit by them, as it shifts to the borders of 
fields, to rastrojo, where quick-growing shrubs offer cover, and also 
settles among coconut groves, mangos, and other trees and shrubbery 
about houses; it even follows clearings well up into the mountains. In 
the older, settled regions, when lawns are developed, in due course the 
Casca Pardo becomes bolder, so that in Balboa and Panama City it 
runs about freely on open lawns, like the American Robin so wide- 
spread in the settled areas of North America. Its usual mannerisms are 
closely similar to the northern bird, much more so than is the case with 
most others of its genus found in Panama. 

Though this thrush is seen constantly, its true abundance is evident 
only when trees that bear edible berries are in fruit. Then the birds 
come in numbers, often boldly, to feed with other birds. Like the 
American Robin (Turdus migratorius) they eat many earthworms 
where these are abundant, and like related thrushes they decoy readily 
when I have used calls to attract small birds. Their alarm note is a 
rapid repetition of a single syllable, pup, pup, higher in pitch than that 
of the northern robin. A common note, rather low, is a curious drawn- 
out whine that suggests vaguely some of the calls of the Smooth-billed 
Ani. 

When the first brief rains come in the latter half of the dry season, 
perhaps in February, the Casca Pardo begins to sing, at first in low 
tone and hesitantly, but by early March males in full song present a 
chorus at the first hint of dawn that has awakened me pleasantly on 
many tropical mornings. Eisenmann writes that the song resembles 
that of the North American Robin, but tends to be softer and sweeter. 
It sometimes includes twee-oo or chwee-oo phrases, which occasionally 
are uttered alone and suggest a Streaked Saltator. Clucking and cack- 
ing calls also suggest vocalizations of its northern congener, although 
often sounding rather less vigorous. Distinctive is a long-drawn, whin- 


FAMILY TURDIDAE 139 


ing call with a nasal quality syllabized by Eisenmann as kee-weé-vah 
or wee-eé-gua. 

The nest, fairly sturdy, rests on a base of leaves and other dried ma- 
terials that often is loose and untidy. The main cup is of grass and 
fibrous materials, with a hardened inner wall of mud, molded by the 
breast of the female. The bottom is padded lightly with fine rootlets 
and grass, and the outside may be covered with green moss. A usual 
site is ina fork in a tree or shrub, anywhere from 1.5 to 10 m above the 
ground (though, rarely, they may be placed much higher). Many lo- 
cations are chosen, from fronds of a palm or a banana, an epiphyte, or 
even the broad leaves of an herbaceous plant, sometimes where support 
is insufficient so that the nest may fall. It is common to find one located 
in some roof angle on a house. For years there were nests on the tops 
of the pillars at the entrance of the old Tivoli Guest House in Ancon. 

The eggs, usually two or three, occasionally four, in form are oval, 
and in color are white with a tinge of green, deeper in some, liberally 
dotted over the entire surface with cinnamon and rufous-brown mark- 
ings, which may appear purple or lilac where thinly covered by chalky 
shell. Some strongly marked examples have a heavy overlay of color 
forming an irregular cap over the larger end. In three sets collected in 
the Canal Zone by Major General G. Ralph Meyer, external measure- 
ments are as follows: 

GZ260)< 202; 28. x 2110) 29'0'x 21.2) and) 29:6 20.5 mma. 
C2 area) Wl9.5 25) 19'5)\and 29/0 x 19,9 1mm. 
(9) 20.9<71.0)and 281x211 mm: 

The principal nesting season is from March to July, with occasional 
records in February and August. In the breeding season of 1970 
Morton (Science, vol. 171, 1971, p. 920) observed 56 nests in Summit 
Gardens, Canal Zone, and found that increased nest predation during 
the rainy season, which began in late April, reduced nesting success 
from the 42 percent of the dry season to 15 percent. He suggested that 
the robins begin breeding in the dry season when food for nestlings is 
scarcer because the increased predation of the rainy season outweighed 
any increase in productivity that might result from greater food avail- 
ability. 

Fully grown immature birds, to be recognized from adults by the 
prominent clay-colored tips on the feathers of the greater and middle 
coverts, are common by the first week in September. Adults are in 
molt at the end of August. After the breeding season, the birds may 
gather at sunset in small companies to roost in groves of trees, often 
near houses. 


I40 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


While known widely as one of the species called casca, locally this 
familiar bird has other names, as platanera in Herrera, capasucia in 
southern Veraguas, and si quiere near Almirante, the last perhaps in 
imitation of its curious whining call. 

The species is widely distributed through Central America north to 
southern and eastern Mexico, (even casually reaching Texas) and 
south to northern Colombia. Northward, the race casius extends from 
Panama through Costa Rica on both slopes. On the Caribbean side it 
crosses the Colombian border in eastern San Blas to Acandi on the 
western shore of the Gulf of Uraba. 

H. Loftin (Eisenmann in litt.) reports one banded at Ancon, Canal 
Zone, on September 6, 1964, recaptured dying on April 13, 1974, at 
Balboa, less than 1 km from the banding site. Another, banded at 
Gamboa on October 23, 1964, was found dead in June 1968 within 3 
km of the banding site. 


TURDUS PLEBEJUS PLEBEJUS Cabanis: Mountain Robin, 
Casca Serrana 


Turdus plebejus Cabanis, Journ. fur Orn., vol. 8, September 1860 (January 1861), 
p.323:; (Costa JRicas) 


Adults brownish gray, darker above, lighter below; with black or 
blackish brown bill. 

Description.—Length 235-250 mm. Adult (sexes alike), above dark 
olive-brown to brownish gray with the crown faintly darker; lighter, 
grayer below, with the abdomen more or less grayish white; throat 
faintly streaked with brownish black; undertail coverts with wedge- 
shaped dusky markings; underwing coverts dull brown, margined with 
tawny. 

Immature, like adult, but with small tips of buff on the outer greater 
primary wing coverts and breast. (These are soon lost, so that the 
juvenile individual rapidly becomes like the adult). 

Kennard (Proc: Boston Soc. Nat: Hist., vol. 38, 1928s) 460) ane! 
male taken March 13, 1926, on the Boquete Trail, noted that the bill 
was “blackish brown; iris raw umber, tarsus brown.” 

Measurements.—Males (11 from Chiriqui), wing 127.7-140.6 
(136.7), tail 92.3-99.9 (94.8), culmen from base 22.7-25.0 (23.7), 
tarsus 31.5-34.5 (33.4) mm. 

Iemales (11 from Chiriqui), wing 122.9-134.8 (130.0), tail 88.0- 
94.5 (92.6, average of 10), culmen from base 22.7-24.4 (23.7), tarsus 
32.1-35.0 (33.5) mm. 


FAMILY TURDIDAE IAI 


Resident. Found in the Subtropical Zone in the mountains of Chir1- 
qui, where it is recorded from Cerro Picacho, on the main volcano from 
near Cerro Punta on the western side, and from above Boquete on the 
east; reported by Griscom from Cerro Flores in eastern Chiriqui. 
Ridgely reports it down to 1000 m in central Chiriqui (Fortuna Dam 
site). Found also on the Caribbean slope, where it was taken by Ken- 
nard at about 1400 m on the Boquete Trail. The species occurs from 
southern Mexico to western Panama; the nominate race in the moun- 
tains of Costa Rica and Panama. 

Griscom’s report of this species from the mountains of Veraguas 
(Bull. Mus. Comp. Zool., vol. 78, 1935, p. 361) appears to be in error, 
as the bird is not known to range eastward beyond the limits given 
above. 

This is a species of the mountain forests, common where found, but 
of restricted range in Panama. | have seen individuals frequently about 
feeding trees, where they often descend to the ground to pick up fallen 
berries in addition to those taken among the branches. They decoy 
readily at a faint squeak, and come near to remain completely motion- 
less, sometimes on open limbs, and sometimes under cover of leaves. 
Occasionally, I have noted them on high perches in dead trees in clear- 
ings. They are usually in small flocks of as many as 15 or 20. 

The cacking or clucking call note is much like that of the North 
American Robin (Turdus migratorius). Kennard and Peters (Proc. 
Boston Soc. Nat. Hist., vol. 38, 1928, p. 460) record the song as more 
musical than that of the northern bird, but Ridgely (1976, p. 275) calls 
the song “inferior to that of its allies, being faster, more repititious and 
monotonous, with few pitch changes.” In addition to the cacking and 
kicking calls, they give a whining call resembling that of the Clay- 
colored Robin, but more musical and more mournful, like od0000reee, 
lasting about 1% seconds, then repeated after an interval of 4-5 
seconds. 

Skutch (Publ. Nuttall Orn. Club, no. 7, 1967, p. 112) found a cup- 
shaped nest hidden among dead leaves and other debris in the crotch 
of a tree, elevated 3 m above the ground. In it were 2 young with 
feathers sprouting. Later, when grown, “the plumage closely resem- 
bled that of [the] parents, except for some rather inconspicuous buffy 
spots or bars on its breast and wing coverts.’”’ No description of the 
eggs has been seen. 

For the date of Cabanis’s description of this thrush, see statement at 
the end of the account of Turdus nigrescens. 


142 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


TURDUS NIGRESCENS Cabanis: Sooty Robin, Casca Tiznada 
Turdus nigrescens Cabanis, Journ. fur Ornith., vol. 8, no. 47, “September, 1860,” 
published January 1861, p. 234. (Volcan Irazu, Costa Rica.) 

Large; adult blackish brown; bill and feet yellow. 

Description.—Length 235-270 mm. Adult male, dull grayish brown 
to blackish brown; wings and underwing coverts dull black; lores and 
area around eye black; throat indistinctly lined with black. 

Adult female, similar to male in general, but duller; throat faintly 
paler, with black lines more distinct. 

Immature, averaging lighter on dorsal surface; crown, back, and 
scapulars streaked with dull buff; greater and middle wing coverts 
tipped narrowly with dull buff to form somewhat indistinct bars; under- 
surface basally dull ochraceous-buff, with the individual feathers tipped 
and barred with dull black; dull buff of throat only lightly marked; 
undertail coverts blacker, narrowly lined with buff; underwing coverts 
dull blackish brown, barred and edged with dull cinnamon-buff. 

In an adult female, taken March 1, 1965, at 2300 m on the west face 
of Volcan de Chiriqui, the iris was ivory-white; bill bright yellow, 
with a faint clouding of dull brown on the nasal operculum, and on the 
side of the culmen adjacent; tarsus and toes bright yellow; claws black; 
bare, thickened outer rim of eyelid honey yellow. Some specimens of 
either sex have the entire bill plain yellow. 

Measurements.—Males (10 from 1900 to 3100 m on Volcan de 
Chiriqui), wing 137.5-144.9 (140.9), tail 107.3-119.4 (114.2), culmen 
from base 23.6-26.1 (24.9), tarsus 37.0-39.3 (37.8) mm. 

Females (10 from 1920 to 3100 m on Volcan de Chiriqui), wing 
134.8-138.9 (137.6), tail 107.2-114.2 (110.4), culmen from base 23.5- 
25.0 (24.4), tail 35.1-38.4 (36.5) mm. 

A series from Costa Rica agrees in size with the measurements 
above. 

Resident on the higher levels of Volcan de Chiriqui, Chiriqui, at 
1800 to 3000 m, also occurs lower on the Volcan de Chiriqui massif, and 
in the high mountains of Costa Rica. 

The little that is known of these mountain thrushes indicates them 
usually to be tame, ranging among trees and also on the ground, alone 
or in scattered company of a few individuals, frequently in pastures 
and clearings. The species, described from Volcan Irazt in Costa 
Rica, was first recorded from Panama by Salvin and Godman (Biol. 
Centr. Amer., Aves, vol. 1, 1879, plate 1, p. 25) from several specimens 
sent to them by Enrique Arcé marked “Volcan de Chiriqui.” Bangs 
(Proc. New England Zool. Club, vol. 3, 1902, p. 49) recorded a series 


FAMILY TURDIDAE 143 


taken by W. W. Brown in May and June 1901 from the high slopes of 
the volcano. Blake (Fieldiana: Zool., vol. 36, no. 5, 1958, pp. 549-550) 
listed 8 in the Monniche collection from elevations of 1980 to 3078 m. 
The specimens examined from Chiriqui appear identical in size and 
coloration with those from the mountains of Costa Rica. 

Eisenmann writes that in western Chiriqui it is known from above 
timberline on Volcan de Chiriqui, but is also frequently seen on the 
ground in the cleared area about the top of the trail between Cerro 
Punta and Boquete, at about 2400 m. On July 21, 1964, at least a 
hundred were in groups of as many as 20 in one small tree at about 
1860 m. It is likely that some downward wandering occurs after the 
breeding season; some of the birds seen were in juvenal plumage. 

Ridgely (1976, p. 276) describes the song as “poor but still some- 
what robinlike” and renders the calls as “trrrr and a harsh tchweerp, 
tchweerp.” We adds (1m litt.) that the Sooty Robin does not seem to 
sing very often. 

A nest found in Costa Rica at Villa Mills, Cartago Province, April 
21, 1974, by Maxine Kiff held two eggs containing feathered embryos. 
The eggs were “robin’s egg blue’ and unspotted. One, now in the 
collection of the Western Foundation of Vertebrate Zoology, measured 
33.00 X 22.42 mm. The nest was located 4 m high in a 7-m tree at the 
edge of cutover montane rain forest and pasture, about 1 m from the 
trunk in the crotch of a limb. The nest was composed of mosses and 
sticks, lined with fine grasses. A nest found by Carriker (Ann. Car- 
negie Mus., vol. 6, 1910, p. 738-739) at Volcan Irazi was made of 
grass and mud. 

The original description of this thrush, in the Journal fur Ornithol- 
ogie, was published in the final issue for the year, which was dated 
September 1860. The copy of this number in the library of the Smith- 
sonian’s Division of Birds when received had a notice printed on the 
bottom of the front cover stating that it had been published “im Januar, 
1861.” This data is carried in C. W. Richmond’s card catalogue for 
nigrescens, and was so listed by Ridgway (Bull. U.S. Nat. Mus., vol. 
50, Part IV, 1907, p. 124). 


HYLOCICHLA MUSTELINA (Gmelin): Wood Thrush, 
Zorzal Pechimanchado 


T.(urdus) mustelinus Gmelin, 1789, Syst. Nat., 1 (2), p. 817, ex Latham “Tawny 
Thrush” (“in Noveboraco” = New York). 


Medium size; upper surface rufescent brown; undersurface white, 
heavily spotted with black. 


144 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Description.—Length 163-192 mm. Sexes alike; undersurface white, 
heavily spotted with black on breast and sides; top of head and hind- 
neck tawny brown to russet, changing to cinnamon-brown and russet 
on back and wings; rump and tail light olive; a distinct white ring 
around eye; side of head dull black, lined narrowly with white; bill 
black, with base of lower part yellowish buff; tarsus and toes flesh 
color. 

Measurements.—Males (10 from North Carolina to Pennsylvania), 
wing 102.4-109.6 (106.2), tail 62.5-72.4 (68.3), culmen from base 
18.9-21.5 (20.2), tarsus 29.7-32.4 (30.9) mm. 

Females (10 from North Carolina to Pennsylvania), wing 102.0- 
106.2 (105.0), tail 65.3-70.0 (67.5), culmen from base) 1992214 
(19.8), tarsus 28.4-32.0 (30.0) mm. | 

Winter resident, found locally October to April, mainly from the 
Canal Zone westward. Panama is at the southern end of this species’ 
winter range, which begins in Texas. There is one record for Co- 
lombia, a male collected west of the Rio Atrato in extreme northwest- 
ern Choco on December 6, 1975; the specimen is in the collection of 
Inderena, Bogota (J. V. Rodriguez, Lozania, no. 31, April 30, 1980, 
p. 8). In their winter home Wood Thrushes live in the undergrowth 
on and above the forest floor in wooded areas where their presence 
may be indicated by their sharp call notes, though the birds usually re- 
main hidden. 

One banded October 8, 1963, at Magnolia Gardens, Charleston, 
South Carolina, by T. A. Beckett was captured alive and released by 
Galindo at Almirante, Bocas del Toro, October 26, 1963. Willis (Liv- 
ing Bird, 1966, pp. 198-200) on Barro Colorado Island found them in 
small numbers from October to April associated in feeding on and near 
the ground with ant swarms. Galindo, in netting and banding opera- 
tions at Almirante, in 1962 recorded them as arriving October 16. They 
have been found with some regularity east to the Canal Zone, but only 
casually beyond. I caught 1 in a mist net at the old Tacarcuna Village 
site in Darién on March 10, 1964, and recorded 2 calling at Mandinga 
in western San Blas, February 7, 1957. 

Ridgely has only once heard the Wood Thrush singing in Panama, 
on March 20, 1979, at Nueva Suiza, Chiriqui. 


CATHARUS USTULATUS (Nuttall): Swainson’s Thrush, 
Zorzal de Swainson 


Medium size; upper surface olive-brown, undersurface white, with 
olive-brown streaks on side of throat and spots on upper breast. 


FAMILY TURDIDAE I45 


Description.—Length 148-185 mm. Sexes alike; upper surface from 
crown to lower back uniform olive to sepia, upper tail coverts slightly 
brighter; wings and tail olive to sepia, with basal third of all but outer- 
most primary and innermost secondary lighter (more distinctly on un- 
dersurface of wing); distinct buffy white streak at lores and eye-ring; 
side of face like back; throat buffy white, streaked with dark brown on 
sides; upper breast washed with pale buff and spotted with dark brown; 
rest of undersurface white with sides color of back; underwing coverts 
buffy. 

The Swainson’s Thrush is the commonest of the North American 
thrushes that migrate through or winter in Panama. It breeds across 
much of northern North America and winters from the southern 
United States to Brazil and Argentina. In extensive netting opera- 
tions at Almirante, Bocas del Toro, Galindo and Méndez ( Bird-Band- 
ing, 1965, p. 234) found during the fall of 1963 that this species formed 
76 percent of all migrant thrushes netted. They captured 3120 Swain- 
son’s, compared with 743 Gray-cheeked Thrushes, 145 Veeries, and 95 
Wood Thrushes. Galindo and Méndez, and Willis (Living Bird, 1966, 
p. 201), who spent the fall of 1960 on Barro Colorado Island, found 
Swainson’s Thrushes commonest in October, ‘‘scattered” or “rare” in 
winter, and uncommon in spring. Griscom (Bull. Mus. Comp. Zool., 
vol. 78, 1935, p. 361) calls it “casual” in summer, based on a specimen 
taken on July 4 at Permé, San Blas. The earliest fall specimen in the 
Smithsonian collection is from October 10, 1960, taken at Almirante, 
Bocas del Toro, and the latest spring specimen was collected on the Rio 
Jaqué, Darién on April 11, 1947. 

Ripley (Checklist Birds World, vol. 10, 1964, pp. 171-173) lists 4 
races of the Swainson’s Thrush. They are analyzed in detail by Bond 
(Proc. U. S. N. M., vol. 114, 1963, pp. 373-387). Specimens of all 4 
have been collected in Panama. C. u. swainsoni is the form that regu- 
larly occurs here; it may be found in wooded and shrubby areas 
throughout the Republic from lowlands and foothills, up to at least 
1800 m in the Chiriqui highlands, and on the Pearl Islands. 


CATHARUS USTULATUS ALMAE (Oberholser) 
Hylocichla ustulata almae Oberholser, 1898, Auk, 15, p. 304. (East Humboldt Mts., 
opposite Franklin Lake, Nevada.) 
Characters —Upper surface distinctly grayer than in other forms. 
Measurements.—Males (10 from Alaska, Mackenzie, and Alberta), 
wing 96.1-102.1 (99.8), tail 66.1-70.6 (68.8), culmen from base 14.2- 
17.7 (16.4), tarsus 27.0-28.7 (27.9) mm. 


146 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Females (10 from Alaska, Mackenzie, and Alberta), wing 93.2- 
100.0 (97.1), tail 60.2-68.2 (62.3), culmen from base 13.5-16.7 (15.7), 
tarsus 26.1-29.3 (27.4) mm. 

This race breeds in southern and eastern Alaska east to western 
Mackenzie and south through the mountains to Colorado and northwest 
Utah. The winter range is unclear, in large part, perhaps owing to the 
difficulty of identifying specimens out of the breeding season. Russell 
(A. O. U. Ornith. Mon. no. 1, 1964, p. 142) does not list this form for 
British Honduras, and Monroe (A. O. U. Ornith. Mon. no. 7, 1968, 
p. 307) considers it “too weakly differentiated to recognize taxonomic- 
ally,” so gives no information for Honduras. Dickey and van Rossem 
(Field Mus. Nat. Hist., Zool. Ser., vol. 23, pub. 406, 1938, p. 457), 
however, call it a “locally common spring migrant” in El Salvador. 
C. u. almae is not mentioned by Slud (Bull. Amer. Mus. Hist., vol. 128, 
1964, p. 299) as occurring in Costa Rica, although Bond (Proc. U.S. 
Nat. Mus., vol. 114, no. 3471, 1963, p. 380) lists 1 collected at Bonilla, 
Costa Rica, April 13, 1908. Miller (Univ. Calif. Pub. in Zool., vol. 66, 
no. 1, 1962, p. 42) collected 1 he believed was C. wu. almae in the western 
Andes of Colombia at latitude 3.5°N on December 26, 1958. 

Nine specimens from Panama in the Smithsonian collection have 
been identified as C. u. almae. One of these was collected in winter, on 
January 20, 1965, at Isla Cébaco, Veraguas; others were taken be- 
tween March 7 and April 6: in Darién at Tacarcuna Village (March 7, 
1964) and Jaqué (April 6, 1946), at Cerro Chame, Province of Pan- 
ama (March 27, 1955), El Barrero, Herrera (March 22, 1948), and 
Isla Brincanco (March 23, 1962) and El Volcan (March 27, 1954), 
Chiriqui. | 


CATHARUS USTULATUS USTULATUS (Nuttall) 


Turdus ustulatus Nuttall, 1840, Man. Orn. U.S. and Canada, ed. 2, pp. vi, 400, 830. 
(forests of the Oregon = Fort Vancouver, Washington.) 


Characters.—Upper surface sepia brown, brightest on rump; more 
rufescent in autumn. 

Measurements.—Males (10 from British Columbia and Washing- 
ton), wing 95.6-102.0 (99.4), tail 66.1-74.6 (71.3), culmen from base 
15.8-17.8 (17.2), tarsus 27.9-31.6 (29.0) mm. 

Females (10 from British Columbia and Washington), wing 92.7- 
98.6 (95.4), tail 65.1-70.8 (67.7), culmen from base 16.5-18.3 (17.1), 
tarsus 27.7-31.4 (29.1) mm. 

This race breeds in southeastern coastal Alaska from Juneau south, 
coastal British Columbia, and west of the Cascade Mountains in Wash- 


FAMILY TURDIDAE WAT, 


ington and Oregon. It migrates through California to winter primarily 
in western Mexico. There are 2 specimens of nominate ustulatus from 
Panama in the Smithsonian collection, from Fl Volcan, Chiriqui, col- 


lected March 17, 1965, and February 26, 1960. 


CATHARUS USTULATUS OEDICUS (Oberholser) 


Hylochichla ustulata oedica Oberholser, 1899, Auk, 16, p. 23. (Santa Barbara, 
California. ) 


Characters.—Upper surface grayer and paler than that of nominate 
ustulatus and buffer, less olivaceous, than swainsoni. 

Measurements.—Males (10 from California, Oregon, and Wash- 
ington), wing 92.0-100.8 (95.2), tail 66.1-74.5 (71.9), culmen from 
base 112.9213.9 (13:5), tarsus 25.3-29.1 (27.7) mm. 

Females (8 from California and Washington), wing 89.0-96.5 
(93.1), tail 64.8-74.7 (69.9), culmen from base 12.7-14.9 (13.5), tar- 
sus 26.7-29.9 (28.1) mm. 

This race breeds in California, southwest Oregon, and along the 
east slopes of the Cascade range to northern Washington. It winters 
from Arizona south to southern Mexico. I have not found any refer- 
ences in the literature to its occurence farther south, but 1 specimen 
in the Smithsonian collection, USNM no. 457921, seems best assigned 
to this race. It is a male collected by Frank A. Hartman at El Volcan, 
Chiriqui, on March 2, 1953. 


CATHARUS USTULATUS SWAINSONI (Tschudi ) 


Turdus Swaimsoni Tschudi, 1845, Faun. Peru., Aves, p. 28; new name for Merula 
zwitlsont Swainson, nec Turdus wilsonti Bonaparte (Carlton House, Saskatche- 
wan.) 


Characters.—Upper surface olive-brown, distinctly less gray than 
C. u. almae. 

Measurements.—Males (10 from Labrador, Nova Scotia, and 
Maine), wing 98.1-102.6 (100.7), tail 66.2-72.2 (68.3), culmen from 
base 15.6-17.6 (17.0), tarsus 26.5-92.3 (27.9) mm. 

Females (10 from Labrador, Quebec, Manitoba, Alberta, Maine, 
and Montana), wing 93.2-103.0 (97.9), tail 64.5-70.0 (67.2), culmen 
from base 15.9-17.5 (16.7), tarsus 27.0-28.8 (27.8) mm. 

This race breeds from eastern Alberta to the Atlantic Coast, through 
Saskatchewan, Manitoba, Ontario, Quebec, and southern Labrador 
south to northern New England and in the Appalachians from New 


148 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


York to West Virginia, and farther west from northern Michigan to 
northern Minnesota. It winters from Mexico to northern Argentina. 

As on its northern breeding grounds, in Panama this is the most 
arboreal of the spotted woodland thrushes. It is a regular visitor to 
fruiting trees and bushes, and also sallies after insects. Willis (Liv- 
ing Bird, 1966, p. 201-203) on Barro Colorado Island found that it 
descends to the ground to follow swarms of army ants, pecking at the 
insects flushed by the ants, and usually driving away Gray-cheeked 
Thrushes and Veeries, but foraging only in niches not occupied by 
resident species. Willis noted that when following army ants in No- 
vember, Swainson’s Thrushes would occasionally deliver fragments of 
song, in addition to call notes. Skutch (A Naturalist in Costa Rica, 
1971, p. 103) found them singing in Costa Rica from March to early 
May. In Panama, Ridgely (1 litt.) has heard in March and April “a 
full whisper song similar to what one hears on migration in eastern 
WISss 

An interesting example of how rapidly these birds can put on weight 
to increase the fat supply needed for migration is recorded by Leck 
(Bird-Banding, 1975, p. 201), who netted a Swainson’s Thrush at the 
Volcan de Chiriqui Field Station of the Florida Audubon Society on 
April 28, 1968, when it weighed 25 g, and retrapped it there 6 days 
later, on May 4, when its weight was 31.6 g. 


CATHARUS MINIMUS MINIMUS (Lafresnaye): Gray-cheeked Thrush; 
Zorzal Carigris 


Turdus minimus Lafresnaye, Rev. Zool., vol. II, no. 1, January, 1848, p. 5. (Bogota, 
Colombia. ) 


Medium size; upper surface dull olive; upper breast streaked irreg- 
ularly with dusky, rest of undersurface white. 

Description—Length 157-187 mm. Sexes alike; upper surface from 
crown to back, olive to grayish olive; tail somewhat browner; side of 
head grayish olive; no distinct eye-ring; ear region streaked narrowly 
with dull white; side of jaw buffy white; undersurfaces of body white, 
with upper breast washed with pale buff; sides of throat to upper breast 
pale buff streaked irregularly with dusky; rest of undersurface white, 
with the sides dull gray; underwing coverts dull gray, edged lightly 
with white. 

In a male collected at Independence, Kansas, May 6, 1905, the iris 
was brown, maxilla and tip of mandible black, rest of the bill dull yel- 
low; tarsus dull brown. 

Measurements.—Males (10 from Alaska to Newfoundland), wing 


FAMILY TURDIDAE I49 


100.0-106.4 (103.2), tail 68.1-75.2 (70.2), culmen from base 15.5-17.9 
GIG) tarsus) 27.1-30.2 (29.5)! mm. 

Females (10 from Alaska to Newfoundland), wing 95.4-99.2 
(97.8), tail 65.2-68.6 (66.4), culmen from base 16.0-17.9 (17.0), tarsus 
27.4-32.2 (29.9) mm. 

Migrant. Gray-cheeked Thrushes are noted especially in the period 
of fall migration; most of those seen are probably the nominate sub- 
species, although bicknelli and aliciae (if it can be distinguished) may 
also occur. Specimens of the nominate race definitely identified are 2 
recorded by Chapman (Auk, 1931, p. 121) taken by R. R. Benson at 
Cocoplum, Bocas del Toro, October 31 and November 1, 1927. Others 
that I have examined are as follows: a male in the British Museum 
taken by Arcé on the Volcan de Chiriqui in 1870 (date not specified) ; 
a male taken by B. Feinstein at the old Tacarcuna Village in Darién 
February 11, 1959; another male from the same locality that I col- 
lected March 5, 1964; 2 (from the Gorgas Memorial Laboratory) 
taken at Almirante, Bocas del Toro, a female on October 12, 1962, and 
another with sex not marked on October 6, 1962. This race winters 
from Nicaragua to northern Peru and northwestern Brazil. 

In extensive netting operations, Dr. Pedro Galindo, assisted by E. 
Méndez and Abdiel Adames ( Bird-Banding, vol. 34, 1963, pp. 202-209) 
banded 264 of these thrushes at Almirante from October 15 to the end 
of November, 1962. None were taken in the following spring period. 
In 1963 the first record for fall was October 2. In 1964, on January 29, 
1 was netted followed by others on April 22 and 24. In fall, the first 
appeared September 25 and the last on November 25. Willis (Living 
Bird, 1966, p. 203) found them fairly common on Barro Colorado Is- 
land in 1960 from October 5 to December 3. He recorded 15 on No- 
vember 1. The following year he noted migrants commonly from 
October 12 to November 24, and single birds seen subsequently on 
January 25, February 4, and April 15, 1961. They were noted regu- 
larly around moving ant swarms, apparently in search of other insects, 
not the ants. 

Loftin (Carib. Journ. Sci., vol. 3, no. 1, 1963, p. 6) reported banding 
65 of this species near Almirante October 19-21, 1962, with 2 on No- 
vember 10-11, 1962, and a few in the Canal Zone near Corozal and 
Curundu in November 1962. 


CATHARUS FUSCESCENS (Stephens): Veery, Zorzal Lomo Oscuro 


Size medium; upper surface dull rufous-brown; sides grayish white, 
without markings. 


150 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


Description—Length, 151-175 mm. Sexes alike; above dull grayish 
brown to somewhat tawny brown; center of throat, lower breast, and 
abdomen white (changing on lower foreneck and upper breast to pale 
butf), streaked and spotted rather lightly with tawny-brown; sides 
grayish white. 

Migrant and winter visitor from the north. Fairly common, found 
from September to early May. 

The Veery, like the Swainson’s Thrush, ranges in forest areas; it has 
been encountered mainly through individuals captured in mist nets. 
Usually they are taken in company with the much more abundant 
Swainson’s Thrush. Of the 4 geographic races currently recognized 
on their northern breeding grounds, 3 are represented in collections 
now at hand from Panama, as indicated under the names that follow. 
As the remaining group, Catharus fuscescens subpallidus Burleigh 
and Duvall (Proc. Biol. Soc. Washington, vol. 72, 1959, p. 33), is m1- 
grant from its breeding grounds like the others, it seems probable that 
it will be identified in the Republic as further data accumulate. It is 
similar in general to C. f. salicicola, differing in duller, more grayish, 
less rufescent coloration on the dorsal surface. 

Sightings cannot be identified as to subspecies and even in the hand 
racial identification can be difficult. Most Panama observations are of 
mist-netted birds on the Caribbean coastal area. G. V. N. Powell and 
S. G. Martin netted many at Ft. Sherman, Canal Zone, between Sep- 
tember 28 and October 22, 1966; the numbers caught were highest be- 
tween October 3 (16) and October 15 (8). None were caught during 
banding done on Cerro Campana. At Almirante, Bocas del Toro, this 
species was taken rather infrequently, although in 1963 and 1964 some 
were taken from the last week in September (earliest Sept. 23, 1963) 
through the third week in November (1 bird), and none in December, 
January, or February. In spring, far fewer birds are netted at Al- 
mirante. 

On the Pacific slope, one was banded by V. Kleen at Nueva Suiza, 
Chiriqui, on October 12, 1967, and Ridgely observed 1 at Ft. Clayton, 
Canal Zone, on September 28, 1968. 


CATHARUS FUSCESCENS FUSCESCENS (Stephens) 
Turdus fuscescens Stephens, in Shaw, Gen. Zool., vol. X, pt. 1, September 1817, 
p. 182. (Pennsylvania. ) 
Characters.—Brighter, more reddish brown, on the entire dorsal 
surface from crown to rump. 


FAMILY TURDIDAE P52 


M easurements.—Males (10 from the breeding range, taken in June), 
wing 98.1-105.1 (101.7), tail 70.0-77.5 (73.7), culmen from base 15.6- 
Passe.) tarsus 29:0-31,0) (29.9) nam. 

Females (10 from the breeding range, taken in May and June), wing 
92.2-100.4 (96.9), tail 64.2-73.4 (67.6), culmen from base 16.2-18.4 
(17.1), tarsus 27.7-30.0 (28.6) mm. 

Migrant from the north, recorded definitely in Panama from 2 male 
specimens: one, in the National Museum, received from the Gorgas 
Memorial Laboratory, collected at Almirante, Bocas del Toro, October 
25, 1963; and a male, in the museum at the University of Cincinnati, 
collected October 10, 1931, at Puerto Obaldia, San Blas, by Hasso von 
Wedel. 

The northern nesting grounds extend from southwestern Quebec 
and southern Ontario south to northeastern Ohio, eastern Pennsyl- 
vania, central New Jersey, and the District of Columbia (Rock Creek 
Park) and the mountains of eastern Tennessee and northwestern Geor- 
gia. In its winter home it has been recorded from Colombia and Ven- 
ezuela to south-central Brazil. The few reports from Panama indicate 
that it is a passage migrant. 


CATHARUS FUSCESCENS FULIGINOSUS (Howe) 


Hylocichla fuscescens fuliginosa Howe, Auk, vol. 18, No. 3, July 1900, p. 271. 
(Codroy, Newfoundland. ) 


Characters —Rather bright reddish brown above; in general, like 
C. f. fuscescens, but decidedly darker. 

Measurements.—Males (10 from Newfoundland and New Jersey), 
wing 94.0-104.6 (100.3), tail 71.4-76.1 (73.3), culmen from base 15.0- 
18.6 (16.2), tarsus 27.8-31.6 (29.3) mm. 

Females (6 from Newfoundland, Rhode Island, and Mississippi), 
wing 94.2-100.2 (96.4), tail 64.2-71.2 (69.0), culmen from base 14.8- 
17.3 (16.4), tarsus 27.9-29.9 (28.9) mm. 

This is the breeding form of Newfoundland, the Magdalen Islands, 
and south-central Quebec. In the north it is recorded as a migrant near 
the Atlantic Coast from Massachusetts to Virginia, and is believed to 
winter in northern South America. 

There are 2 specimens at present known from Panama, both taken 
during Galindo’s banding studies at Almirante, Bocas del Toro, a male 
on October 25, 1962, and a female in October 1963 (without more def- 
inite date). 


152 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


CATHARUS FUSCESCENS SALICICOLA (Ridgway) 


Hylocichla fuscescens salicicola Ridgway, Proc. U.S. Nat. Mus., vol. 4, no. 240, 
April 13, 1882, p. 374. (Fort Garland, Colorado.) 


Duller, less reddish brown, averaging darker above as compared to 
fuscescens or fuliginosa. 

Measurements.—Males (10 from Manitoba, Montana, Colorado, 
and Wisconsin), wing 95.4-102.0 (99.4), tail 66.3-79.3 (72.5, average 
of 9), culmen from base 14.5-18.5 (16.7), tarsus 27.7-31.5 (30.3) mm. 

Females (9 from Minnesota, North Dakota, lowa, Montana, and 
Colorado), wing 90.0-101.3 (94.8), tail 65.0-73.4 (70.0), culmen from 
base 14.6-17.4 (15.9, average of 8), tarsus 27.0-32.3 (29.1) mm. 

Breeds from southern British Columbia, Alberta, Manitoba, Nevada, 
Colorado, South Dakota, and Minnesota to southern Wisconsin, Indi- 
ana, and southwestern Ohio; recorded in winter in South America from 
Colombia, Venezuela, and Mato Grosso, Brazil. 

Two specimens have been taken at Almirante, Bocas del Toro, dur- 
ing banding studies: a female October 12, 1962, and a male marked 
October, without more certain date, 1963. 


CATHARUS MEXICANUS FUMOSUS Ridgway: Black-headed 
Nightingale-Thrush, Zorzal Cabecinegro 


Catharus fumosus Ridgway, Proc. U.S. Nat. Mus., vol. 10, August 6, 1888, p. 505. 
(Costa Rica.) 


Size medium; crown in male dull black, in female brownish black. 

Description.—Length 146-164 mm. Male, crown black; upper sur- 
face, including tail brownish olive; band across upper breast and lower 
foreneck, sides, underwing coverts and feathered area on upper legs 
somewhat brownish olive; throat, upper foreneck, breast, and undertail 
coverts white; wings like tail, but with outer webs of primaries red- 
dish brown. 

Female, similar but crown brownish black. 

Juvenile (male, Rio Changuena, September 8, 1961), crown, back, 
and wing coverts blackish brown, with feathers lined centrally with 
dark buff; upper breast buff centrally, feathers edged and tipped with 
dusky black; lower breast and abdomen dull buff. 

Measurements.—Males (10 from Veraguas and Costa Rica), wing 
81.3-88.5 (84.5), tail 54.8-64.5 (60.3), culmen from base 14.1-17.5 
(15.8), tarsus 28.4-32.7 (30.8) mm. 


Iemales (10 from Panama, Costa Rica, and Nicaragua), wing 78.5- 


FAMILY TURDIDAE 153 


87.0) (82.8), tail 52.1-61.6 (57.5), culmen from base 13.5-17.5 (15.5, 
average 01/9) tarsus 28:2-31.8 (30.1) mm. 

Resident. Found locally in the higher elevations on the Caribbean 
slope of the mountains in western Panama. Recorded to date from the 
Boquete trail, and the upper Rio Changuena, 1060 m, both in the high- 
lands of western Bocas del Toro; Calovévora, and the Cordillera del 
Chuct, Veraguas (around Santa Fé, high on both slopes). Beyond 
Panama this race is recorded north to Nicaragua; other races are 
found north into Mexico. 

In the original description, Ridgway listed his type as “Adult male 
(No. 101765, Costa Rica, October 20, 1881; José C. Zeledon).” Zele- 
don, in his Catalogo de las Aves de Costa Rica (Anal. Mus. Nac., 1887, 
published 1888), cited under each species the localities and number of 
specimens in the collection of the Museo Nacional. On page 104, family 
Turdidae, he wrote “Catharus fumosus Ridgway, Jiménez 1,’ this 
being a specimen in the local collection (not Ridgway’s type, which is 
in the Smithsonian collections). Deignan (Bull. U.S. National Mu- 
seum, 1961, vol. 221, pp. 429-430), with reference to Zeledon, above, 
listed Ridgway’s type as “ “Costa Rica’ = Jiménez, Province of Limon, 
Costa Rica,” but no such specific locality can be ascribed to the actual 
type. 

Slud (Birds of Costa Rica, 1964, pp. 300-301) speaks of fumosus as 
a species of the Caribbean slope in the humid foothills of that country, 
found frequently in the same habitat as its relative Catharus fuscater 
hellmayri, low down in undergrowth, often on a log or on the ground, 
remaining rather constantly undercover, moving with quick hops, then 
coming to a sudden halt. 

In November 1940, on the watershed leading to the Caribbean slope 
on Cerro Santa Maria, Costa Rica, I found these thrushes fairly com- 
mon, ranging near the ground in the dense shadows of the humid forest. 
When detected they disappeared immediately in the heavy cover. At 
this season they were quiet, and were not singing. 

Slud (op. cit.) says the song is variable, with one or two short 
phrases, with an extra note or two added at the start or end. In some 
phrases there is a slurred, sharp “‘sreek.”’ The quality is fuzzy, slurred, 
or skewy flutelike; some notes are wavering warbles. Calls are a quick 
dry trill, a rattle like stones knocked together, a very soft unclear 
pseer, and an antbirdlike grrr and meww. C. Hartshorne character- 
izes the song as “barely mediocre,” weaker and inferior to that of C. 
frantzu. 


154 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Carriker (1910; Ann. Carn. Mus. vol. 6, p. 749) reports 2 nests on 
the lower foothills north of the Volcan de Turrialba, Costa Rica, at 
about 600 m, that on April 20 each held 2 eggs. The eggs were pinkish 
white, very thickly and finely speckled with reddish brown over the 
entire surface; average measurements were 24X18 mm. The nests 
were almost entirely of green moss with a lining of skeletonized leaves. 
Each nest was placed about 1.6 m from the ground in a small palm in 
heavy forest cover. 


CATHARUS FUSCATER (Lafresnaye): Slaty-backed Nightingale-Thrush, 
Zorzal Pizarreno 


FicureE 12 


Upper surface dark slate color; side of head and chin deep black; 
upper foreneck and throat white. 

Description—Length 160-180 mm. Adult (sexes alike), above, 
dark slate, the head somewhat blacker, wings and wing coverts slightly 
grayer; breast, foreneck, and sides gray; center of breast and abdomen 
white. 


FicureE 12.—Slaty-backed Nightingale-thrush, Zorzal Pizarrefio, Catharus 
fuscater. 


This wide-ranging species is found in mountain areas from Costa 
Rica and Panama to the Andes of northwestern South America, where 
it ranges from Colombia and northwestern Venezuela to northern Bo- 
livia. Three subspecies are recognized in Panama. 


FAMILY TURDIDAE 155 


CATHARUS FUSCATER HELLMAYRI Berlepsch 
Catharus fuscater hellmayri Berlepsch, Orn. Monatsber., vol. 10, no. 5, May 1902, 

p. 69. (Chiriqui.) 

Characters.—Gray of undersurface, including sides and flanks, defi- 
nitely darker; throat grayer. 

A juvenile in the Museum of Comparative Zoology from Coliblanco 
de Sarapiqui, on Irazu, Costa Rica, taken August 23, 1899, has the 
entire crown fuscous-black; hindneck natal brown; back, rump, scapu- 

lars, and wing coverts between natal brown and bone brown, wings and 

tail fuscous; throat and upper foreneck buffy brown, with the bases of 
the feathers grayish white; sides of neck, breast, and area adjacent 
olive-brown, with indistinct spots of dull chamois, and basal shaft 
streaks of grayish white; flanks and undertail coverts olive-brown. 
The culmen (in the dried skin) is fuscous; rest of the bill, tarsi, and 
feet light brownish yellow, indicating that these areas were light- 
colored in life, as they are in the adult bird. 

An adult male in the Museum of Comparative Zoology collected 
above Boquete, Chiriqui, March 1, 1901, has the following color notes 
on the label, made by W. W. Brown, Jr., “Iris white; orbital ring and 
tarsus orange-vermilion; upper mandible vermilion, region of nostrils, 
and tip of culmen dusky; under mandible vermilion.”’ 

Measurements.—Males (10 from Chiriqui), wing 88.0-94.4 (89.7), 
tail 64.6-73.6 (68.5), culmen from base 17.5-19.0 (18.2), tarsus 34.1- 
36.8 (35.4) mm. 

Females (4 from Chiriqui), wing 78.6-81.9 (89.7), tail 54.0-60.4 
(see), culmen trom base 15.3-17.8 (16.5); tarsus 31.3-35.0 (33.4) 
mm. 

Resident. Found in the mountains of Chiriqui, recorded on the 
eastern face of the volcano above Boquete at 1200-3000 m, and in 
eastern Chiriqui at Chame, and above Tole, including the upper valley 
of the Rio Caldera (Bajo Mono). 

This race was known early in Panama, mainly from a series collected 
in 1901 by W. W. Brown, Jr., on the mountain slopes above Boquete 
(sans Iroc) New nel Zool) Club, vol! 3; 1902) p. 50). It may be 
restricted in distribution, as I did not find it during several seasons on 
the western and northern slopes of Volcan de Chiriqui, from above 
I] Hato and Cerro Punta to near Sereno on the Costa Rican boundary, 
nor on the headwaters of the Rio Chiriqui Viejo. From eastern Chiri- 
qui, Salvin (Proc. Zool. Soc. London, 1867, p. 132) received an early 
specimen taken by Arcé in the Cordillera de Tolé, above Tole. 

Ridgely (in litt.) suggests that the restricted range may be based on 


150 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


the fact that this species occurs only in humid areas, while most of 
Volcan de Chiriqui, especially the west side, is relatively dry. The bird 
is common in wetter areas to the east, such as the Fortuna Dam site, 
where Ridgely found up to 5 birds at ant swarms in February-March 
1976: 

Older statements of range for hellmayri, in addition to Chiriqui, 
have included Veraguas, based apparently on the report by Salvin 
(Proc. Zool. Soc. London, 1870, p. 180) of specimens collected by 
Arcé from Calovévora, and Cordillera del Chucu, in the mountains 
north of Santa Fé. The report requires confirmation 1f the skins, 
which may be 1n the British Museum, can be located. 

Slud (Birds of Costa Rica, 1964, p. 301), describes the song of 
hellmayri as clear and flutelike in tone, with a definite pattern in its 
syllables. Ridgely described the songs of birds at Fortuna as “tooo teee, 
tee-too-too-tee, with some variation, beautiful clear whistles.” 

M. A. Carriker (Ann. Carnegie Mus. vol. 2, 1910, p. 748) found a 
nest in Costa Rica at Juan Vinas, May 20, 1907, with two fresh eggs. 
“The nest is not so pretty a structure as that of mexicanus, being con- 
structed of leaves, moss, and weedstems, and lined with fine weed-fibres 
and grass. It was placed in a low bush in the heavy forest. The eggs 
are pale blue, thickly speckled and dotted and blotched over the entire 
surface with light chestnut-rufous. Measurements: 25X18 mm.” 


CATHARUS FUSCATER MIRABILIS Nelson 


Catharus fuscater mirabilis Nelson, Smiths. Misc. Coll., vol. 60, no. 3, September 
24 (September 27), 1912, p. 24. (Head of Rio Limon, 1585 m elevation, Cerro 
Pirre, Darién.) 

Characters.—Similar to Catharus f. fuscater, but gray of foreneck 
and upper breast duller; throat grayish white; white of lower breast 
and abdomen duller; dorsal surface slightly paler. 

Measurements.—Males (6 specimens), wing 83.3-88.4 (86.6), tail 
68.0-70.9 (69.5), culmen from base 17.5-18.7 (18.3), tarsus 33.2-37.5 
(34.6) mm. 

Females (4 specimens), wing 78.8-82.9 (80.7), tail 57.2-64.0 (60.9), 
culmen from base 17.6-19.5 (18.4), tarsus 32.7-34.7 (33.5) mm. 

Resident. Found in the Subtropical Zone on Cerro Pirre, Darién, 
at 1500-1600 m; specimens in the Gorgas Memorial Laboratory were 
taken at Cerro Cana (August 1965) and Alturas de Nique (April 
1972). 


E. A. Goldman, who collected the series from which this race was 


FAMILY TURDIDAE 157 


described, found it fairly common in undergrowth. He described the 
song as three short syllables given in sequence, with the middle one 
slightly higher in tone than the others. In late April they were in pairs 
and were breeding. A set of two eggs that he collected on April 30, 
1912, are very pale greenish white, spotted lightly with dots of cin- 
namon-brown, which are grouped to form a faintly-indicated cap on 
the summit of the blunt end. They measure as follows: 24.7 17.7, and 
25.0 18.3 mm. 

In both Darién populations the distinct yellow wash on the under- 
' parts of fresh birds disappears in a relatively short time (fide P. Ga- 
lindo; also noted by Eisenmann). 


CATHARUS FUSCATER FUSCATER (Lafresnaye) 


Myioturdus fuscater Lafresnaye, Rev. Zool., vol. 8, September 1845, p. 341. (Bo- 
gota, Colombia.) 


Characters.—Upper foreneck and throat white; side of head and 
chin deep black. 

An adult male, taken February 24, 1964, on Cerro Mali, Darién, 
had the iris pale brownish white; thickened border of eyelid yellow; 
space above nostril, and distal end of culmen dull black; rest of bill, 
tongue, and inside of mouth bright orange; gape orange-yellow; tarsus 
and toes orange-yellow; claws yellowish brown. A female, collected 2 
days later, was similar to the male. 

Measurements.—Males (10 from Cerro Tacarcuna and Cerro Mali, 
Darién), wing 85.4-90.7 (87.9), tail 69.4-76.0 (72.6), culmen from 
base 17.1-19.5 (18.4), tarsus 33.8-35.3 (34.4) mm. 

Females (9 from Cerro Tacarcuna and Cerro Mali, Darién), wing 
81.7-86.8 (83.3), tail 61.4-70.5 (66.8), culmen from base 17.6-19.6 
(18.4, average of eight), tarsus 32.3-35.0 (33.6) mm. 

Resident. Subtropical Zone on Cerro Tacarcuna and Cerro Mali, 
Darién, 600 to 1500 m; fairly common. 

Found locally, low in open undergrowth on the slopes of the moun- 
tains, in part in cloud forest. In late February (1964), gonads were in 
resting stage. In life, with the birds perched quietly near at hand, the 
bright colors of eye, bill, and feet are prominent. 

This race appears to range through the mountains to Ecuador and 
the eastern Andes of Colombia, and to the Sierra de Perija in north- 
western Venezuela. In Panama, it extends across the higher slopes of 
Tacarcuna. Nothing is recorded of its nesting. Specimens in the Amer- 
ican Museum of Natural History collected by Anthony on the eastern 


158 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


base of Tacarcuna in Colombia are similar to those seen from Panama. 

Some form of C. fuscater occurs in the mountainous Cerro Azul- 
Cerro Jefe area of eastern Province of Panama, not far east of the 
Canal Zone. E. S. Morton writes of its song there, heard on May 29, 
1966, “‘a clear series of whistles like a solitaire in quality but slightly 
similar to a Wood Thrush in phrasing. A minor tone—each series of 
three whistles followed by two whistles, each whistle a single pure tone: 
tu tee tu (last note a half tone lower than first) followed by tu tee; then 
a longer pause (3 seconds), and it starts all over again. It was an ex- 
ceedingly beautiful sound coming out of the mist on Cerro Azul.” E. O. 
Willis, present at the time, considered the song pleasing, but rather 
simple, in fact, much simpler than that of C. frantzi. Ridgely, on 
Cerro Jefe on July 24, 1975, heard one singing with the same rhythm, 
but he syllabized it as to-to-tee, tee-too. 

Willis believes that similarity in song suggests that C. fuscater may 
be a representative of the spotted C. dryas, which is absent from south- 
ern Middle America, but reappears in South America. Eisenmann 
feels the evanescent yellowish tones and vague grayish to olive mottling 
below also suggest the relationship. 


CATHARUS FRANTZII WETMOREI Phillips: Ruddy-capped Nightingale- 
Thrush, Zorzal Cabecirrojizo 


Catharus frantzu wetmorei Phillips, Auk, vol. 86, no. 4, October 30, 1969, p. 615. 
(Boquete, Chiriqui.) 


Rather small; upper surface brown; throat and abdomen white, rest 
of undersurface gray. 

Description.—Length 145-170 mm. Adult. Male, crown russet- 
brown; back, rump, and wing coverts somewhat duller; outer webs of 
wings grayer; side of head grayish olive, with area in front of eye in- 
definitely black; throat dull white, becoming gray streaked with white 
centrally on upper foreneck; lower foreneck and upper breast dull gray, 
changing to lighter gray on breast and sides, and to white on lower 
breast, abdomen and undertail coverts; undersurface of wings dull 
dark gray; tibia dull dark gray. 

Female, somewhat paler russet-brown above. 

A male, taken at 2300 m on the north slope of Volcan de Chiriqui, 
Chiriqui, February 24, 1965, had the iris dark brown; maxilla black; 
mandible yellowish orange, becoming brighter orange along the com- 
missure, yellower elsewhere; inside of mouth, including the tongue, 


FAMILY TURDIDAE 159 


orange; tarsus and toes grayish brown, becoming paler brownish white 
along the narrow thin knifelike posterior margin; claws of two lateral 
anterior toes dull brownish white; others grayish brown. 

An adult, taken on the same date, was like the male except that the 
tarsus and toes were somewhat duller colored. 

Measurements.—Males (10 from Chiriqui), wing 84.7-90.5 (87.3), 
tail 66.0-73.1 (70.2), culmen from base 17.6-19.5 (18.4), tarsus 33.3- 
36.7 (34.5) mm. 

Females (10 from Volcan de Chiriqui and Cerro Punta, Chiriqut), 
~ wing 78.3-84.2 (81.2), tail 62.0-68.1 (64.0), culmen from base 17.6- 
18.8 (18.0), tarsus 32.1-34.4 (33.2) mm. 

Resident on the high slopes of the volcano in western Chiriqui, 
mainly above 2000 m, but fairly common down to 1600 m at Cerro 
Punta, found also in southern Costa Rica; other races range from 
northern Costa Rica through Mexico. Phillips (Auk, 1969, pp. 605- 
623) reviews the characters separating this species from C. occidentalis. 

These birds range over the mountain slopes in undergrowth and 
usually remain concealed. They move about quietly and are difficult 
to observe unless one appears in the bushes bordering a trail, on the 
ground at the edge of a pasture, or on the banks of a stream. In late 
February and early March, they occurred in pairs, and were not espe- 
cially shy. 

Occasionally one came beneath the window of our quarters, where 
it hopped quickly over the ground or rested quietly, with head erect. 
The call was a low kree-ee-eet, rather harsh in sound. They responded 
quickly to an imitation of it. 

There has been no account to date of the nest and eggs of this bird 
in Panama. Carriker (Ann. Carnegie Mus., vol. 6, 1910, p. 747) in an 
account of the closely allied race, Catharus frantzu frantziu, on Vol- 
can Irazu in Costa Rica, found 2 nests, April 13 and 14, 1902, one of 
which he described as “made entirely of green moss, but lined with 
fine grass and rootlets. It is very large and bulky for the size of the 
bird, but the cavity of the nest is small... [the nest was] placed on sprays 
of bamboo hanging over the side of a deep ravine, and about seven feet 
from the ground. The eggs are pale blue, thickly speckled and blotched 
with cinnamon-rufous and lilac, thickest about the larger end, in one 
ege forming a cap of rufous and lilac. Measurements: 24.5 to 25.5x 
NSto 19mm.” 

In the Bambito-Cerro Punta area of western Chiriqui, this species, 
although shy and hard to see, is common, usually on or near the ground, 


160 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


in and bordering forest from about 1500 to 2250 m. The song is liquid 
and long-continued, far-ranging, but not nearly so loud as those of the 
local solitaire. One of the frequent phrases can be syllabized as dleédle- 
edleéyee. The song is reminiscent of that of the northern Hermit 
Thrush, the effect melancholy, the quality liquid, but rather throaty; the 
phrases are deliberate with pauses of 2 to 3 seconds between them, and 
the song so long continued that over a hundred phrases may be given 
before there is a rest of more than a very few seconds. Dr. C. Hart- 
shorne, im litt. to Eisenmann, characterized the song very well as hav- 
ing phrase structure close to that of the northern Wood Thrush, but 
tone quality closer to that of the Hermit Thrush. 


CATHARUS AURANTIIROSTRIS (Hartlaub): Orange-billed Nightingale- 
Thrush, Zorzal Pico Anaranjado 


Turdus auranturostris Hartlaub, Rev. Mag. Zool., March 1850, p. 158. (Caracas, 
Venezuela. ) 


Rather small; crown gray; rest of dorsal surface rufescent brown, 
breast gray, rest of undersurface white. 

Description.—Length 132-158 mm. Adult (sexes alike), crown 
gray; rest of dorsal surface, including wings and tail, rufescent brown; 
upper breast and sides gray; throat white, streaked with gray; lower 
breast, abdomen, and undertail coverts white. 

Immature, back and, to a lesser degree, the crown dull dark brown 
with the feathers tipped irregularly with black; breast and sides dull, 
faintly buffy white, with the feathers tipped and edged with dull black; 
undertail coverts buffy brown; bill black. 

Widely distributed as a species from northern Mexico through 
Central America and northern South America, in Colombia, Venezuela, 
and the island of Trinidad. Several forms are recognized, two of them 
found in Panama. (For notes on this species see Zimmer, Auk, 1944, 
pp. 404-408.) The gray-crowned populations of southwestern Costa 
Rica, Panama, and western Colombia were formerly regarded as a 
separate species, C. griseice ps. 

Ridgely (1976, p. 278) describes the song as “poor and unmusical 
with varied phrases, some twangy, others squeaky; a short tsip, wee-ee, 
tsirrip-tsip is one of several.” 

This species tends to range lower in the highlands of Panama than 
the other species of Catharus and is the most widely distributed, favor- 
ing clearings and second growth. 


FAMILY TURDIDAE 161 


CATHARUS AURANTIIROSTRIS RUSSATUS Griscom 
Catharus griseiceps russatus Griscom, Amer. Mus. Nov., no. 141, October 31, 1924, 

p. 6. (Boruca, Costa Rica.) 

Characters——Upper surface darker. Crown darker gray; back, 
wings, and tail duller, darker reddish brown; gray of undersurface 
darker. 

In a male taken on Volcan de Chiriqui, February 2, 1954, the bill, 
including a ring around the nostril and the base of the culmen, was 
light brick red; line of culmen otherwise dark neutral gray; eyelids 
light brick red; iris dark brown; tarsus and toes dark honey yellow; 
claws buffy brown. 

A female, on March 13, had the ring around the nostril, the sides of 
maxilla and mandible, and a ring around the eye bright orange; rest of 
maxilla dark neutral gray; iris dark brown; tarsus and toes yellowish 
orange, with a line down the front of the tarsus, the central portion of 
the toe scutes, and the toes fuscous. In another female, March 19, 1965, 
the iris was light brown; thickened eyelid orange; culmen and sides of 
mandible fuscous-brown; rest of bill, gape, and entire inside of mouth, 
including the tongue, reddish orange; tarsus and toes deep yellow, with 
one or two toes on each foot tinged with brown. 

Measurements.—Males (9 from Chiriqui), wing 76.7-82.9 (79.4), 
tail 51.3-61.4 (57.4), culmen from base 15.5-18.4 (17.1), tarsus 29.4- 
a7 (30.8) mm. 

Females (7 from Chiriqui and Costa Rica) wing 72.3-78.3 (75.4), 
tail 54.3-58.1 (56.6), culmen from base 15.3-18.4 (17.3), tarsus 28.9- 
es (0:3 mm. 

Resident in the mountainous area of western Chiriqui from the 
Costa Rican boundary eastward, mainly from 1000 to 1650 m on the 
slopes of Volcan de Chiriqui. 

Ridgely found this species in north-central Chiriqui on the upper 
Rio Chiriqui (Fortuna Dam site) in a clearing at 1000 m on March 3, 
1976, but, without specimens, it cannot be determined to what race the 
birds belong. 

Eisenmann regards auranturostris as a poor, unmusical singer. The 
song is variable and often rapid. He has syllabized it at Bambito, on 
the western slope of the volcanic massif, as ts-teyt, teweet, tisteet; 
witsteeyt, steeweea, tiststeet; and tsipeeareet, tseea; elsewhere he has 
heard tsip, wee-ee, tsirrip-tsip. Usually only the second phrase can be 
called musical. Sometimes there are twanging wang or snip notes in- 
terposed. Songs heard about Boquete were generally of similar quality. 


162 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Found in low, dense thickets in high forest on Silla de Cerro Pando, 
they were always secretive and difficult to see. Occasionally, I heard 
them give low, mewing, sometimes trilling calls. One (prepared as a 
specimen) was captured lower down in a mist net set near the Rio 
Chiriqui Viejo. Two birds in juvenile dress, fully grown, were taken 
by S. L. Olson and K. Blum at Bambito, at about 1600 m, July 3, 1966. 

Blake (Condor, 1956, p. 388) described a nest in the Monniche col- 
lection from Lérida, Chiriqui, as located 2.5 m above the ground in a 
small tree, a compact structure of moss, much like that of the Black- 
faced Solitaire, but with a few bits of dried leaves and plant stalks 
woven into its base and sides. The cup, neatly lined with black hairlike 
rootlets, had outside dimensions of 544x414 x3 inches, with the cup 
22X13 inches. The single egg preserved, pale greenish white, thickly 
and almost uniformly speckled with light reddish brown, measured 
23.4X17.3 mm. The collector’s field notes state that the second egg, 
similar in color, measured 23 X18 mm. 


CATHARUS AURANTIIROSTRIS GRISEICEPS Salvin 
Catharus griseiceps Salvin, Proc. Zool. Soc. London, May 1866, p. 68. ( Veraguas. ) 


Characters.—Upper surface paler; crown grayer; back lighter 
brown; breast and sides lighter gray. 

A female collected at El Copé, on the Pacific slope of the Province of 
Coclé had the iris dark brown; edges of eyelids and bill orange, with 
the culmen dark neutral gray; tarsus and toes light orange; claws light 
brown. 

Measurements.—Males (10 from Chiriqui and Veraguas), wing 
72.5-82.5 (78.3), tail 55.5-62.0 (55.8, average of 9), culmen from base 
15.1-17.8 (1615); tarsus 28/5-31.3 (30.2) mama: 

Females (9 from Chiriqui, Veraguas, and Coclé), wing 73.5-82.5, 
tail 54.6-62.5 (59.5), culmen from base 13.1-18.6 (16.1), tarsus 29.5- 
3730/6) mam: 

Resident. Found locally in Chiriqui (Chame, Barriles, San Félix), 
Veraguas (Sona, Sante Fé) and Coclé (El Copé). 

Near San Félix I found them in a small tract of forest near the Rio 
San Félix, and near the coast in the area called Macano, also in forest. 
At El Copé, on February 25, 1962, in the hills on the Pacific side, the 
one taken was in heavy undergrowth in a small tract of forest. 

Eisenmann (in litt.) saw the species singing above Santa Fé, Vera- 
guas, at approximately 750 m, near the Agricultural School, on March 
28, 1974. 


FAMILY TURDIDAE 163 


CATHARUS GRACILIROSTRIS Salvin: Black-billed Nightingale-Thrush, 
Zorzal Piquinegro 


Catharus gracilirostris Salvin, 1865, Proc. Zool. Soc. London, (1864), p. 580. 

(Volcan de Cartago, Costa Rica.) 

Rather small; bill black. 

Description.—Length 140-155 mm. Adult (sexes alike), head and 
neck slate-gray, becoming paler, whiter, on the throat; crown dark 
brownish gray, lighter, grayer on the forecrown; back of head, hind- 
neck, back, and rump somewhat reddish brown; wings and tail duller 
but with brighter edgings; undersurface plain gray changing to white 
on the abdomen; a broad band of pale dull brown across the upper 
breast; throat lined narrowly and indistinctly with white; underwing 
coverts dark gray. 

Immature, upper surface as adult; throat gray; breast, sides, and 
flanks reddish brown; increasingly buffy on center of belly; abdomen 
white. 

Found in Chiriqui on the high slopes of the volcano above Boquete, 
and on the west face; recorded also in eastern Chiriqui on “Cerro 
Flores” (Cerro Santiago), and in the western end of the Serrania de 
Tabasara (near boundary between Chiriqui and Bocas del Toro). The 
nominate race 1s found in the mountains of Costa Rica. 

Little is available as yet on the habits of this interesting bird. In the 
original description of the race accentor, Bangs included the observa- 
tion by Brown that, like its relatives in the genus, it “is a fine song- 
ster.” Brown’s series, 8 adults of both sexes, was collected on the 
higher slopes of the volcano from about 1500 to 3000 m elevation. 
Griscom found it in eastern Chiriqui near the western end of the Ser- 
rania de Tabasara at about 1800 m, where he collected 2 (male and 
female), in March 1924. With limited comparative material he de- 
scribed these as a separate race bensoni, which later he found not valid. 
However, a specimen collected by R. H. Pine on the Chiriqui-Bocas del 
Toro border 23 km north-northeast of San Félix on June 16, 1980, 
matches Griscom’s original description of bensoni and is easily dis- 
tinguishable from the series of specimens from western Chiriqui in the 
Smithsonian collections. 

Eisenmann writes that on the Volcan de Chiriqui massif in the area 
above Cerro Punta it is the highest ranging nightingale-thrush, usually 
the commonest above 2100 m on the trail to Boquete. It seems less shy 
than its allies, often feeding on the ground in cleared patches. On a 
recently plowed field at about 2250 m, Eisenmann saw one member of 
a feeding pair chase away one of a pair of the larger C. frantzii, but the 


164 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


bird returned. Generally, this species is seen on or a few feet above the 
ground. The song suggests in pattern that of C. frantzi, but in Eisen- 
mann’s opinion, much inferior, being weaker, with a squeaky quality. 
The nest and eggs are little known. Carriker (Ann. Carnegie Mus., 
vol. 6, 1910, p. 745) described the egg of C. g. gracilirostris, the closely 
similar subspecies of Costa Rica found on Volcan Irazu, as “blue, more 
or less thickly speckled and dotted with reddish brown or deep chest- 
nut, usually with a heavier wreath or cap of this color at the larger end, 
though rarely speckled evenly over the entire surface. Measurements: 
20.5 to 23X15 to 16 mm.” 


CATHARUS GRACILIROSTRIS ACCENTOR Bangs 


Catharus gracilirostris accentor Bangs, Proc. New England Zool. Club, vol. 3, 
January 30, 1902, p. 50. (Volcan de Chiriqui, 10,200 ft. (3140 m), Chiriqut, 
Panama. ) 


Characters.—Forehead gray, rest of upper surface light reddish 
brown; band across breast paler than in bensoni. 

An adult male, caught at Volcan de Chiriqui, March 4, 1965, had 
the iris dark reddish brown; thickened edge of eyelid fuscous-black; 
bill black, with the tip of maxilla and of mandible on the inner side 
dark slate; rest of inside of mouth, including tongue and gape, orange; 
tarsus dull brown, paler on back on the plantar edge; claws fuscous- 
black. 

Measurements.—Males (8 from Chiriqui), wing 71.2-78.4 (75.4), 
tail 56.7-66.7 (61.9), culmen from base 15.1-17.0 (15.8, average of 7), 
tarsus 30.9-33.4 (32.0) mm. 

Females (6 from Chiriqui), wing 70.8-74.8 (73.1), tail 54.6-62.2 
(59.3), culmen from base 14.3-18.3 (15.8), tarsus 30.0-31.1 (30.5) 
mm. Hp 

Resident. Known from the highlands of western Chiriqui. In the 
Smithsonian collections there are 2 males taken March 4 and 9, 1965, 
to 2650 and 2700 m on the west face of Volcan de Chiriqui. A male and 
female were collected also near Cerro Punta, March 12, 1962, by C. L. 
Hayward. Blake (Fieldiana Zool., vol. 36, no. 5, 1958, p. 552) listed 
4 males and 4 females in the Monniche collection from western Chiriqui 
from Casita Alta, Pefia Blanca, and Volcan de Chiriqui from 2040 to 
3130 m, without other comment than the localities with date and eleva- 
tion. (For this series he used the name Catharus gracilirostris gracili- 
rostris. ) 


FAMILY PTILOGONATIDAE I 65 


CATHARUS GRACILIROSTRIS BENSONI Griscom 


Catharus gracilirostris bensoni Griscom, Amer. Mus. Nov., no. 141, October 31, 
1924 p. 7. (Cerro Flores, 6000 ft, eastern Chiriqui, Panama.) 


Characters.—Forehead blackish brown; rest of upper surface dark 
reddish brown; band across breast dark reddish brown. 

Measurements.—Male (1 from Cerro Flores, Chiriqui, the type 
specimen), wing 68.9, tail 62.8, culmen from base 15.3, tarsus 30.5 mm. 

Females (2 from eastern Chiriqui), wing 68.0-69.0 (68.5), tail 59.7- 
62.7 (61.2), culmen from base 14.2-15.0 (14.6), tarsus 29.7-31.8 
(30.8) mm. 

Resident. Known from the highlands of eastern Chiriqui, where 
in 1927 Benson collected a male and female for Griscom at 2000 m in 
Cerro Flores and where R. H. Pine collected two at 1800 m on Cerro 
Bollo, 23 km north-northeast of San Félix. 

[This race was described by Griscom from a small series taken on 
Cerro Flores in easternmost Chiriqui. Most authors have not recog- 
nized it and it was considered to be a synonym of accentor in the origi- 
nal draft of this manuscript. Subsequently, however, R. H. Pine 
collected 2 specimens on the Chiriqui-Bocas del Toro border 23 km 
north-northeast of San Félix, Chiriqui, on June 16, 1980, that necessi- 
tate a reappraisal of this form. These specimens are from very near the 
type locality of bensomi and were preserved in alcohol, from which one 
was prepared as a skin. This specimen is much darker and more rufous 
dorsally than accentor and has a much darker, fuscous crown. The 
colors do not appear to have been affected by alcohol, which if anything 
usually causes fading and lightening in color. The fresh specimen 
matches Griscom’s original description of bensoni exactly, whereas the 
type series of bensont, although still distinguishable from accentor, 
shows somewhat less pronounced differences, suggesting that they may 
have been affected by aging. In any case, from the evidence available 
now it would appear that the race bensoni is quite distinct and should 
pe maintamed. S/O] 


Family PTILOGONATIDAE: Solitaires and Silky-flycatchers, 
Solitarios y Dorales Sedosos 


These attractive birds form an assemblage that may or may not be 
related. The solitaires usually are placed in the Turdidae, but Sibley 
(Auk, 1973, p. 394-410), after a review of past analyses and an exami- 


166 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


nation of egg-white proteins, concludes that they are more closely re- 
lated to the silky-flycatchers and should be placed in that family, in a 
position next to the Turdidae. The silky-flycatchers, sometimes given 
subfamilial rank in the Bombycillidae, were found by Sibley to be less 
similar to the waxwings than to the solitaires. Ames (Bonn. Zool. 
Beitr., vol. 26, 1975, pp. 125-127), in an examination of ¢syameeal 
morphology in the Muscicapidae, concludes that the generalized oscine 
syrinx in Myadestes places it outside the Turdidae, but not necessarily 
in the Ptilogonatidae. 

The 2 solitaires and 2 silky-flycatchers found in Panama are all high- 
land birds living in forest and forest edges where they feed on fruit 
and insects. All but the Varied Solitaire are restricted to Costa Rica 
and western Panama; this last has an equally narrow distribution in 
the highlands of eastern Panama and extreme northwestern Colombia. 

The solitaires are shy and their drab plumage makes them difficult 
to see in the thick foliage, but their song makes up for their inconspicu- 
ousness. The Black-faced Solitaire’s loud, liquid, seemingly studied 
whistles are among the most beautiful of bird songs. The song of the 
Varied Solitaire, from what little is known, is harsher. Although es- 
sentially arboreal except when they follow foraging bands of ants, 
solitaires build their cup-shaped nest on or close to the ground. 

The silky-flycatchers, in contrast, are strikingly colored in patterns 
of yellow and gray or black. The tapered, elongated tail of Ptilogonys 
caudatus, black with white spots, is flashed dramatically in aerial acro- 
batics when the birds are pursuing insects. Also, unlike the solitaires, 
their voices are undistinguished and their nests are placed in bushes 
and trees. 


KEY TO SPECIES OF PTILOGONATIDAE 


I. Yellow present on. undersurface... 62a). ..05 ose teen) see Z 
No yellow on tinderstirface! 2.5. hoe cin. pole oie ae he ee 5) 
2. Tail long, graduated, with white spots. 
Long-tailed Silky-flycatcher, Ptilogonys caudatus. p. 171 
Tail not graduated, all black. 
Black-and-yellow Phainoptila, Phainoptila melanoxantha, p. 175 
3. Back dark blue-gray. 
Black-faced Solitaire, Myadestes melanops. p. 167 
Back bright reddish brown. 
Varied Solitaire, Myadestes coloratus. p. 169 


FAMILY PTILOGONATIDAE 167 


MYADESTES MELANOPS Salvin: Black-faced Solitaire, 
Solitario Carinegro 


FIGURE 13 
Myiadestes melanops Salvin, Proc. Zool. Soc. London, for 1864, February 1865, 

p. 580, pl. 35. (Tucurrique, Costa Rica.) 

Medium size; body color rather dark gray, with face, wings, and 
tail black. 

Description.—Length 157-186 mm. Adult (sexes alike), body slate 
color throughout, slightly paler on lower surface; side of head, includ- 
ing space around eye, cheeks, forehead, and chin, deep black; wings 
black except for the base of the tertials, outer webs of the secondaries, 
and inner webs of inner primaries, which are edged with gray; tail 
black, except the outer web and distal end of the outer rectrix, which 
are brownish gray; some of the longer rectrices with a small terminal 
spot of white. 

Juvenile, in first plumage, dark sooty black, the feathers with a term1- 
nal spot of light tawny brown; undersurface with the light markings 
more extensive. 

Adult male, iris dark brown, varying individually in depth of color; 
external edge of eyelids narrowly dark brown; bill, tongue, and inside 
of mouth orange; tarsus and toes honey yellow; tips of claws dull neu- 
tral gray. 

M easurements.—Males (10 from Chiriqui), wing 84.5-91.1 (87.9), 
tail 76.1-80.5 (77.5), culmen from base 14.2-16.5 (15.1), tarsus 20.5- 
23.2 (21.4) mm. 

Females (10 from Chiriqui and Bocas del Toro), wing 85.8-91.2 
(89.0), tail 74.1-80.7 (77.3), culmen from base 13.1-16.1 (15.0), tarsus 
ee 25).5) 021.1) mm. 

Resident. Found locally in the higher mountain forests in Chiriqui 
and Bocas del Toro (Rio Changuena, 1975 m, Almirante); on the up- 
per slopes of the Chiriqui volcano from 1200 to 2900 m (fairly com- 
mon above Boquete and the Bambito-Cerro Punta area); in the moun- 
tains on the upper Rio Changuena, northwestern Bocas del Toro, in 
heavy forest, down to 750 m; recorded eastward in the mountains of 
eastern Veraguas (on the Pacific slope at Santa Fé, 830 m, Chitra, 
1170-1500 m; on the upper Caribbean slope near Calovévora and 
Cordillera del Chuct). 

There are a few specimens labeled as from lowland localities near 
sea level. Three collected by Batty from the Pacific Coast islands Brava 
(January 28, 1902) and Cébaco (February 6, 1902) in the American 


168 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Museum of Natural History are probably mislabeled. One taken in a 
mist net at Almirante, Bocas del Toro (October 28, 1963; Gorgas Me- 
morial Laboratory) could have been either a stray or a cage-bird 
escape. 

The solitario is a forest inhabitant, living mainly on tree-grown 
slopes or in thickets in ravines and gullies. Though shy and retiring, 
they may be seen moving under cover in low vegetation, along shaded 
cut banks near roads and streams, or, casually, flying across open trails. 
Their songs may be heard regularly, but country-dwellers assure the 
traveler that it is impossible to see the bird at freedom! In my observa- 
tions, they are not gregarious, as I encountered them singly. They 
were captured rather regularly in mist nets. 


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Ficure 13.—Black-faced Solitaire, Solitario Carinegro, Myadestes melanops. 


In Costa Rica, where they are better known, they are prized as cage 
birds for their songs. Skutch, in his detailed study of the solitaire in 
that country (Publ. Nuttall Orn. Club, no. 7, 1967, p. 117), says that it 
“sings with a calm deliberation that seems the product of a studied art.” 


FAMILY PTILOGONATIDAE 169 


C. B. Worth (Bird-Lore, vol. 41, 1939, p. 283) in late July and early 
August noted that it was not then in song, as he heard it utter only a 
“loud rasping shoo-wee-ee.” Eisenmann writes that their songs in 
Chiriqui are loud, clear, and deliberate, with a liquid quality suggestive 
of the better Catharus singers, but louder, and often with a sharpness 
that reduces the pleasing quality. The phrases are usually separated by 
pauses of about three-fourths of a second. Songs transliterated as 
leedideedleé, tleéah, lee-dah, lee-dee may alternate with songs consist- 
ing of three, two, or one phrase, usually with components like tleedlee, 
- leeee, leydee, lee-dah, leedo, leedoo, leeoo-lee or the like. They sing 
most abundantly in the late afternoon, after 4 p.m. 

Blake (Condor, 1956, pp. 387-388) described a nest in the Monniche 
collection, found March 30, 1932, near the Finca Lérida above Boquete 
“located in the crevice of a rock about three feet above a road. It is 
made almost wholly of soft moss and has a smooth lining of black hair- 
like rootlets. Dimensions: 6X5xX2¥4 inches, the cup 34ZX2Y%UxKY% 
inches. The two eggs, partly incubated, measured 2416.7 and 22.8X 
16.9mm. They are pinkish white, minutely speckled with dull chestnut 
except toward the larger end where they are increasingly spotted and 
blotched with the same.” 

The solitaire is reported to feed on berries, including those of palms, 
other trees and shrubs, and also on insects. In Costa Rica, Slud and 
Skutch have found it ranging near foraging bands of ants, presumably 
attracted by the insects that are disturbed. Two males collected at 
Cerro Punta by Strauch (Bull. Brit. Orn. Club, 1977, p. 64) weighed 
60.0.and 32.6 ¢. 


MYADESTES COLORATUS Nelson: Varied Solitaire, Solitario Variado 


Myadestes coloratus Nelson, Smiths. Misc. Coll., vol. 60, no. 3, September 27, 
1912, p. 23. (Cerro Pirre, 1524 m elevation, Darién.) 


Similar to Myadestes melanops, but with back and wing coverts 
rufous-brown. 

Description.—Length 162-181 mm. Adult (sexes alike), crown, up- 
per hindneck, and entire lower surface gray; back, wing coverts, and 
tertials rusty brown; forehead, throat, space around eye, and anterior 
area of side of head black; tail with three outer rectrices tipped with 
white. 

Juvenile, compared to that of melanops, distinctly brown above, with 
pale markings more extensive, crown more heavily spotted; abdomen 
marked indistinctly with white. 


LO BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Two adult males, collected on Cerro Mali, Darién, February 21, 
1964, had the iris dark brown; edge of eyelid orange-brown; bill, in- 
side of mouth, and tongue orange; tarsus and toes somewhat paler 
orange; claws orange-yellow. 

Measurements.—Males (10 from Cerro Mali and Cerro Pirre), 
wing 86.5-93.2 (88.8), tail 74.2-81.2 (77.5), culmen from base 13.6- 
15.4 (14.6), tarsus 20.5-23.1 (21.9) mm. 

Females (10 from the same areas), wing 84.4-88.7 (86.8), tail 71.2- 
78.8 (75.2), culmen from base 13.2-15.3 (14.2), tarsus 20.8-22.8 
(21.8) mm. 

Resident. Found locally on Cerro Pirre (1500 to 1600 m), Cerro 
Mali (1400 to 1460 m), Cerro Quia (900 m), Aturas de Nique, Darien. 

In the latter part of February 1964, these solitaires were fairly com- 
mon in forest on the slopes of Cerro Tacarcuna, ranging mainly low, 
concealed in the leaves in the tops of low undergrowth. Generally they 
remained hidden in this fairly dense cover, so that most of those taken 
for specimens were captured in mist nets. Rarely, one was seen in the 
forest. Also rarely, we heard a single note of the song, harsher in 
sound than that of melanops of the mountains in Chiriqut. 

No nests have been recorded. Young recently from the nest and fully 
grown birds in juvenile dress were noted in the latter part of May and 
in June. A female (AMNH) from “Mt. Tacarcuna, 4600 ft.” is labeled 
“ovary large” on April 16. A full grown molting juvenile (Gorgas 
Mem. Lab) was taken at Cerro Cana on August 9. 

The species of the genus Myadestes in Panama, obviously of related 
ancestry, form a highly attractive group. Hellmayr (Cat. Birds Amer., 
pt. 7, 1934, pp. 440-443) treated them with those of the Andes as geo- 
graphic races under the name ralloides (described by d’Orbigny from 
Bolivia). All are residents of the intermediate and higher elevations in 
mountain areas, where it may be presumed that their obvious differ- 
ences may have been established during Pleistocene time. In the pres- 
ent Recent period those of Panama divide in two isolated groups, sepa- 
rated in range by considerable spaces of lowland, and here regarded as 
species. 

The western form melanops of the mountains of Costa Rica and 
western Panama, with wholly gray body color, black facial mask and 
yellow bill, is isolated by the broad lowlands of the central isthmus from 
the eastern coloratus group of the mountains of eastern Darién. These 
agree with melanops in black mask and yellow bill, but differ completely 
in wholly brown back. Though this second group is like the forms of 
the Andes in brown back, it is separated geographically from them by 


FAMILY PTILOGONATIDAE yt 


the broad break of the lowlands across northern Choco 1n northwestern 
Colombia. The Andean birds, which lack the facial mask and have 
smaller, dull-colored bills, form the distinct species ralloides, with three 
or more closely similar subspecies ranging south to Peru and Bolivia. 


PTILOGONYS CAUDATUS Cabanis: Long-tailed Silky-flycatcher, 
Capulinero Colilargo 


Ficure 14 

Ptilogonys caudatus Cabanis, 1861, Journ. f. Orn., 8 (1860), p. 402. (Irazu, 
Costa Rica.) 

Medium size; sleek, crested, tail more than half total length of bird; 
blackish wings and tail; body gray and yellowish green (male) or all 
yellowish olive (female). 

Description.—Length 200-244 mm. Male, crown pale gray; bushy 
crest olive-yellow; rest of upper surface slate gray, slightly lighter on 
rump; wing coverts slate gray; primaries and secondaries black, with 
outer webs edged slate gray; tail black, tapered, with central pair of 
feathers elongated, round white spots on outer four pairs; eye-ring 
(sometimes incomplete) lemon yellow; throat pale olive-yellow, be- 
coming more intense on breast; band across chest slate gray; flanks 
yellowish green; abdomen pale yellow; undertail coverts lemon yellow. 

Female, like male but back and wing coverts olive-green, tinged gray 
on rump; undersurface olive-green becoming grayish olive on abdo- 
men. 

Immature, body gray-green; undertail coverts whitish; wings and 
tail as adult. 

One (sex not recorded) taken on Volcan de Chiriqui, Chiriqui, Feb- 
ruary 24, 1965, had the iris reddish brown, bill black, tarsus, toes, claws 
black. 

Measurements.—Males (10 from Chiriqui), wing 91.0-101.0 (95.9), 
tail 120.0-143.3 (130.2), culmen from base 12.3-15.4 (14.1), tarsus 
18.6-22.4 (20.3) mm. 

Females (10 from Chiriqui), wing 84.8-96.0 (90.4), tail 106.7-119.3 
Giits.); culmen from base 12'5-14.9 (13.6), tarsus 17.5-21.7 (19.7) 
mm. 

Resident. Common in the Temperate Zone in highlands of western 
Chiriqui above 1500 m (Ridgely, 1976, p. 281) in forest edges and in 
pastures and clearings with tall trees. Blake (Fieldiana: Zool., vol. 36, 
no. 5, 1958, p. 552), who says the bird “has seldom been reported below 
an altitude of 8000 feet [= 2400 m],” lists specimens collected by Mon- 
niche at Alto de Chiquero, Casita Alta, Copete (summit), Lérida, Pena 


172 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Blanca, and Volcan, at elevations between 1800-3090 m. A male col- 
lected by Hartman at Cerro Copete (2100 m), near Boquete is now in 
the Smithsonian collections, and another male from FE] Salto (1740 m), 
also near Boquete 1s now in the California Academy of Sciences col- 
lection. Bangs (Proc. New England Zool. Club, vol. 3, 1902, p. 58) 
wrote that in May and June 1901, W. W. Brown “found this species 
only at high altitudes [3000-3300 m] on Volcan de Chiriqui. The birds 
were in small flocks, composed of adults and full-grown young still in 
nestling plumage. Sometimes he saw one of these flocks leave the high 
peak of the mountain and fly across the valleys in the direction of the 
lofty Costa Rican mountains, which could be seen in the distance.” 

Ridgely (in htt.) found small numbers at the Fortuna Dam site in 
central Chiriqui, including several down to 1200 m, the lowest elevation 
from which it has been recorded; Fortuna is also the easternmost lo- 
cation from which it is known. 

Besides westernmost Panama, this species is found only in Costa 
Rica, at similar elevations. 

I have encountered this bird several times on Volcan de Chiriqui and 
at Cerro Punta. On February 25, 1955, I took a male in near breeding 
condition at 2010 m in Bajo Grande, Cerro Punta. It was feeding on 
the drupes of a small tree in partly open forest. The tongue was dis- 
tinctly bifid at the tip like that of a titmouse. I saw these birds also on 
Cerro Picacho above 1800 m, but found them so wild that they flew 
before I could shoot. Their flight is swift and direct, but somewhat 
undulating. 

Skutch (Auk, 1965, pp. 375-426) is the only one to have studied the 
breeding biology of this species. From February to July 1963 he ob- 
served several pairs in the trees around a large dairy farm on the 
western end of the massif of Volcan Barba, in the Cordillera Central 
of Costa Rica. Their diet was composed of large quantities of insects 
caught in the air in long sallies and of berries, including those of 
Fuchsia arborescens, Eurya theoides, Citharexylum moccinnu, and a 
mistletoe, all picked while the bird was perched. 

Long-tailed Silky-flycatchers are highly vocal but practically song- 
less. The call most frequently heard is a dry che chip, che chip, given 
both while perched and in flight. Another call given when taking off 
and in flight is a long-drawn, dry che-e-e-e-e. The song, which Skutch 
heard rarely, is composed of “low, lisping notes that were scarcely audi- 
ble at a distance of 25 yards”; it was delivered by the male from a high, 
exposed perch while his mate incubated. 

These birds are loosely social, often traveling in flocks of more than 


FAMILY PTILOGONATIDAE 173 


a dozen that may casually split apart or regroup in flight or at a feeding 
tree. Similarly, nests are often situated within 25 to 100 m of one an- 
other in “colonies” of up to five pairs, or may be entirely by themselves. 


Figure 14.—Long-tailed Silky-flycatcher, Capulinero Colilargo, Ptilogonys 
caudatus, male. 


The birds defend an area around the nest of no more than 30 m, some- 
times less, and often disregard intruders entirely. 
The nesting season, in Costa Rica at least, begins in April, with the 


174. BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


peak of nest building and egg laying in late April and early May. Nests 
are situated from 2 to 20 m off the ground in trees or, rarely, large 
shaded shrubs; the average height of the 19 nests Skutch found was 
about 7 m. The bulky, cup-shaped nest, made almost exclusively of 
beard-lichen (Usnea), with a small amount of cobweb or caterpillar 
silk to bind the lichens together, is placed in the fork of an erect branch 
or in a stout horizontal limb, where upright twigs provide support. 
Males seem to choose the site and begin construction, although they are 
soon joined by their mates, and together are intensely active in bringing 
material to the nest—Skutch watched one pair that made 42 visits in an 
hour. The finished nest has dimensions of 114-127 mm in overall di- 
ameter, 50-64 mm height, 57-64 mm diameter of the cavity, and 41 m 
cavity depth. 3 

Egg laying begins 2 to 4 days after construction is finished; the two 
eggs that form a complete clutch are laid a day apart. Skutch describes 
them as “‘long-ovate and rather pointed, and the shells are without much 
gloss. The ground color is pale gray, of a shade that almost matches 
the bed of lichens on which the eggs lie. On this gray background, 
blotches and spots of shades of dark brown and pale lilac are variously 
distributed. Often there is a distinct belt of heavy brown pigment 
around the thickest part of the egg.’ The measurements of 10 eggs 
averaged 25.8 X 17.2 mm. 

Incubation is done exclusively by the female and lasts 16 or 17 days, 
during which time she is frequently fed by her mate, although she also 
leaves the nest often to feed herself. At hatching the nestlings have a 
distinctive plumage of short, compact tufts of whitish down arranged 
in narrow rows surrounded by large bald areas. By the age of 9 days 
their eyes are open and contour feathers are beginning to expand on 
most parts of the body; in addition, secondary down now covers much 
of the upper surface that was bare at hatching. The young are fed 
several times an hour by each parent. In their first few days of life 
they receive an almost exclusively insect diet; later more fruit is brought 
to them, although they continue to be fed insects as long as they are in 
the nest. At 16 or 17 days of age they begin climbing about beyond the 
edge of the nest, but not until 23-25 days do they fly more than a few 
feet and leave the nest altogether, after which they are still attended for 
several days. By early June they can catch insects by themselves. 

Skutch found that of the 15 nests for which he knew the outcome, 4 
successfully produced fledglings. From the others the eggs or nestlings 
vanished, possibly taken by Blue-throated Toucanets (Aulocorhynchus 
caeruleogularis) or Brown Jays (Psilorhinus morio). 


FAMILY PTILOGONATIDAE 175 


PHAINOPTILA MELANOXANTHA Salvin: Black-and-yellow Phainoptila, 
Capulinero Orinegro 


Phainoptila melanoxantha Salvin, 1877, Proc. Zool. Soc. London, p. 367. (Costa 
Rica = San Francisco, fide Hellmayr, 1935.) 


Phainoptila melanoxantha minor Griscom, 1924, Amer. Mus. Novit., no. 141, p. 8. 
(Cerro Flores, eastern Chiriqui, Panama.) 


Medium size; male mostly shining black with bright yellow on flanks; 
female mostly olive-green with bright yellow on flanks. 

Description.—Length 180-217 mm. Male, entire upper surface ex- 
cept rump shining black; rump lemon yellow; face, throat, and upper 
breast shining black; band across lower breast yellowish green; sides 
and undertail coverts lemon yellow; rest of undersurface gray; under- 
wing coverts yellowish green. 

Female, crown shining black; hindneck bluish gray; back and wing 
coverts yellowish green, with flecks of bluish gray and slightly brighter 
green On rump; primaries, secondaries, and tail feathers brown, edged 
yellowish green; throat gray; band across upper breast olive-yellow; 
sides and undertail coverts lemon yellow; rest of undersurface gray; 
underwing coverts pale yellow. 

Immature male, like female, with a few black feathers on throat. 

Measurements.—Males (10 from Chiriqui, Panama and Irazu, 
Costa Rica), wing 95.4-103.8 (99.5), tail 82.2-90.0 (85.5), culmen from 
base 16.0-19.2 (18.0), tarsus 25.6-28.9 (27.2) mm. 

Females (10 from Chiriqui, Panama, and Irazu, Costa Rica), wing 
92.3-99.0 (96.4), tail 77.4-85.4 (81.0), culmen from base 15.2-17.6 
(16.8), tarsus 25.4-28.8 (26.2) mm. 

Resident. Uncommon in Temperate Zone highlands of western 
Chiriqui and Bocas del Toro (one record, Blake, Fieldiana: Zool., vol. 
36, no. 5, 1958, p. 553) and from eastern Chiriqui and Veraguas. The 
Monniche specimens described by Blake came from elevations of 1560- 
3300 m. Ridgway (U.S. Nat. Mus. Bull., vol. 50, part 3, 1904, p. 124) 
gives the range of elevation as 1200 to 3300 m. Ridgely (in litt.) col- 
lected a female above the Fortuna Dam site, central Chiriqui, at 1150 m, 
on February 27, 1976; the specimen is in the collection of the Gorgas 
Memorial Laboratory. 

Apparently commoner in Costa Rica, this bird was first listed for 
Panama by Salvadori and Festa (Bol. Mus. Zool. Anat. Comp. R. 
Univ. Torino, vol. 14, no. 339, 1899, p. 3) on the basis of a specimen 
collected by Arcé labeled only Chiriqui. Bangs (Proc. New England 
Zool. Club, vol. 3, 1902, p. 58) discusses specimens collected by W. W. 
Brown, Jr., during January-May 1901 from Volcan de Chiriqui and 


176 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Boquete. Blake, cited above, gives several western Chiriqui localities 
as well as the single record from Bocas del Toro on the Caribbean 
slope. In 1924 Griscom reported 4 specimens in extreme eastern Chiri- 
qui, at Cerro Flores (1800 m), nearer the mountains of Veraguas than 
to the Volcan de Chiriqui. He described these as a new race, minor 
(Am. Mus. Nov., no. 141, 1924, p. 8), “similar to the typical form from 
Costa Rica, but averaging smaller; male not differing in color; female 
with hind-neck more extensively gray of a slightly darker shade; rump, 
upper tail-coverts, and edgings to tail feathers slightly greener, less 
yellow.” In 1926 Benson collected at Chitra from 1290 to 1500 m on 
the Pacific slope in Veraguas, the farthest east from which this bird is 
known; Griscom (Bull. Mus. Comp. Zool., vol. 78, 1935, p. 363) as- 
signed these to minor. The race minor has generally not been admitted 
(e.g., Greenway, in Peters, Check-list Birds World, vol. 9, 1960, p. 
373), and its invalidity was confirmed by Pasquier and Olson on exami- 
nation of specimens from both of the series attributed to minor. 

I have never encountered this bird myself, and most of what is 
known of its biology comes from Skutch’s work in Costa Rica (Auk, 
1965, p. 420-422), where in 1963 he observed them near the western 
end of the massif of the extinct Volcan Barba, in the Cordillera Central. 
Here he found them from 2040 to 2250 m, usually within the edge of 
the forest, where the canopy was broken by the removal of a few trees, 
and in adjoining shady pasture. They moved about mainly in the mid- 
dle levels of the tall oaks, flying out from exposed branches to catch 
insects and then continuing on to another branch, unlike many flycatch- 
ing birds which return to their initial perch. Fruit is also an important 
part of their diet; Skutch found them eating the berries of Drimys 
winteri, Boccoma frutescens, Monnina, and Ardisia. They pick the 
fruit either while perched on the plant or while fluttering in front of it. 

Black-and-yellow Silky-flycatchers are far less social than Long- 
tailed Silky-flycatchers, which in fact they hardly resemble. They are 
usually found singly or in pairs, and are less active, never catching fly- 
ing insects with the acrobatic manoeuvers of Ptilogonys. The only calls 
that Skutch heard were “‘a weak chip rather like a wood warbler’s note” 
given by a male and “low, soft notes” from a foraging pair. 

The nest and eggs of Phainoptila were unknown until 1972, when 
they were discovered in Costa Rica by Lloyd Kiff. With F. G. Stiles 
and B. K. MacKay, he found a nest on May 2 at 2400 m on Volcan 
Boas, Alajuela Province; it was 1.5 m from the ground in the central 
crotch of a 2-m sapling, in a thicket so dense as to make the nest in- 


FAMILY SYLVIIDAE EF 


visible from above or the sides. Kiff (in litt.) describes it as “a large 
compact open cup composed mostly of green moss interspersed with a 
few slender stems and fern fronds. It was lined with fine rootlets and 
plant stems. The outer diameter of the nest was 22 X16 cm, and it was 
about 12 cm in depth. The inner cup was 3 cm in diameter and 2 cm 
deep.” Two other nests of Phainoptila from Costa Rica were dis- 
covered in similar situations at the same time of year. 

The 2 eggs from Kiff’s nest were “‘subelliptical in shape and slightly 
glossy. They had a grayish-white ground color with a dense sprinkling 
of fine light gray, purplish brown and dark brown spots over their en- 
tire surfaces. Each egg contained a slightly developed embryo.” They 
measured 27.84X20.30 and 27.19X19.20 mm, with empty dry shell 
weights of 0.274 and 0.260 g, respectively. Kiff concludes that, “aside 
from their somewhat blunter shape,” the eggs of Phainoptila melano- 
xantha “are simply larger versions of those of Phainopepla nitens or 
Ptilogonys cinereus.” 


Family SYLVIIDAE: Gnatwrens and Gnatcatchers, Cazajejenes 


The gnatwrens and gnatcatchers are both usually placed in the family 
Sylviidae, the Old World warblers. Some put the gnatcatchers in a sub- 
family (Polioptilinae) of a broadened Muscicapidae, which also in- 
cludes the subfamily Sylviinae, or in an entirely separate family, the 
Polioptilidae, or reduce them to a tribe (Polioptilini) of the Sylviinae. 
The gnatwrens have in the past been considered to be wrens or even 
antbirds; Rand and Traylor (Auk, 1953, p. 334-337) and Parkes 
(Proc. Biol. Soc. Wash., vol. 93, 1980, pp. 61-67) review the history of 
their classification. Although gnatwrens superficially resemble ant- 
birds, they have an oscine syrinx. Rand and Traylor concluded that 
Ramphocaenus and Microbates are most closely allied to certain Old 
World warblers, especially the African genus Macrosphenus, than to 
Polhoptila. Beecher (Auk, 1953, pp. 279, 328) considers resemblances 
to Old World genera to be convergent. 

Of the 9 species of gnatcatchers, 2 are found in Panama. They share 
the typical gnatcatcher characters of a slim build, gray and white plum- 
age, and a long tail continually wagged or cocked as the bird flits 
through the foliage searching for insects. The gnatwrens are also in- 
sect eaters and, like the gnatcatchers, build cup-shaped nests that often 
are decorated with mosses, leaves, or lichens. 


178 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


KEY TO SPECIES OF SYLVIIDAE 


1.’ Upper surtace brown; billvery lone... 00.2.2. le ee 2 
Upper ‘surface gray, ‘bill relatively‘short.//-... .. oe. 0) es ee 3 
2. Undersurface below throat buffy, tail long. 
Long-billed Gnatwren, Ramphocaenus melanurus rufiventris. p. 183 
Undersurface below throat gray, tail short. 
Tawny-faced Gnatwren, Microbates cinereiventris, p. 186 
3. General coloration light bluish gray. 
Tropical Gnatcatcher, Polioptila plumbea. p. 178 
General coloration dark slaty gray. 
Slate-throated Gnatcatcher, Polioptila schistaceigula. p. 182 


POLIOPTILA PLUMBEA (Gmelin): Tropical Gnatcatcher, 
Cazajején Tropical 


Ficure 15 


Todus plumbeus Gmelin, 1788, Syst. Nat., 1, pt. 1, p. 444. (Surinam.) 


Small, slender, with long tail; upper surface gray and black; under- 
surface light gray and white. 

Description.—Length 92-103 mm. Adult male, crown, hindneck, 
and upper parts of sides of neck black, with a tiny white feather or two 
behind the nostril; back, scapulars, rump, and wing coverts bluish 
gray; primaries and secondaries dusky neutral gray, with outer webs 
edged with slate gray, except for the two outer primaries; tertials edged 
broadly with white on outer webs, the edging becoming pale neutral 
gray toward the tip; tail black centrally, the two outermost rectrices 
white, the next two black at base, the fourth black, tipped white; lores 
and a broad superciliary white; eye-ring white; a conspicous black line 
extending from the eye to the back of the nape; rest of side of head, 
throat, ventral area of sides of neck, abdomen, and undertail coverts 
white; rest of undersurface pale bluish gray; underwing coverts, and 
inner webs of primaries and secondaries, toward the base, white. 

Adult female, like male, but upper surface entirely dark bluish gray, 
without black on the head. 

Juvenile male, like adult female but crown slate-gray and white su- 
perciliary not fully developed behind eye. 

The Tropical Gnatcatcher is a widespread species, occurring from 
southeastern Mexico to central Peru and central Brazil, with habits 
similar to the Blue-gray Gnatcatcher (P. caerulea) of temperate North 
America. In Panama it is found up to about 1200 m almost every- 
where that there are trees, from forest canopy and borders to scrubby 
areas to brushy savannas (Aldrich and Bole, Scient. Pub. Cleveland 


FAMILY SYLVIIDAE 179 


Mus. Nat. Hist., vol. 7, 1937, p. 23). Ridgely notes that it occurs in 
the elfin cloud forest on the wind-swept summit of Cerro Jefe (900- 
1000 m), eastern Province of Panama, where few other lowland birds 
occur. 

This species is an active searcher of foliage and twigs, moving rapidly 
from branch to branch, constantly wagging or raising its tail to dif- 
ferent angles. Usually encountered singly or in pairs, it often joins 
mixed foraging flocks. The diet seems to be purely animal matter: 
Po. Goldman collected 2 at Cana, Darien, on May 26, 1912; the 
‘stomach of one contained 2 pentatonid nymphs 37%, a large fulgorid 
35%, an ant 8%, bit of caterpillar skin 10%, spider remains 10%; the 
other contained a small caterpillar 25%, 2 small scarabaeids 20%, 3 
curculionids (Cryptorhynchini) 15%, a small chrysomelid 5%, a small 
pentatonid 20%, homopteran remains 15%, bits of a spider with 2 or 
more eggs 5%. One taken by Burton near Pacora, Province of Pan- 
ama, weighed 5.9 g (Bull. Brit. Orn. Club, 1975, p. 85); 2 weighed by 
Strauch (Bull. Brit. Orn. Club, 1977, p. 64) were 6.2 and 6.4 g. 


Figure 15.—Tropical Gnatcatcher, Cazajején Tropical, Polioptila plumbea, 
male (below), female (above). 


The vocal repertoire includes “a rather thin but musical song con- 
sisting of a series of simple notes with decreasing intensity, sweet, weet, 
weet, weet, weet, sometimes faster and sibilant; the usual call is a tzeet- 


180 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


tzeet” (Ridgely, 1976, p. 279). Eisenmann (in litt.) has heard nasal 
twanging or mewing notes, sounding like twee, twee, twee and peéoo, 
peéoo, peéoo. 


POLIOPTILA PLUMBEA SUPERCILIARIS Lawrence 


Pohoptila superciliaris Lawrence, 1861, Ann. Lyc. Nat. Hist. New York, 7, p. 304. 
(New Grenada, Isthmus of Panama = Atlantic slope, near Panama Railroad; 
fide Lawrence loc. cit. p. 322.) 


Characters.—Dorsal surface bluish gray; lower foreneck and breast 
whitish. 

A male collected at Las Palmitas, Los Santos, on January 25, 1962, 
had the iris dark reddish brown; base of mandible neutral gray; rest 
of bill black; tarsus and toes dusky neutral gray; claws black. | 

Measurements.—Males (10 from Los Santos, Herrera, Coclé, and 
Province of Panama), wing 43.5-48.6 (45.8), tail 36.2-46.3 (42.1), 
culmen from base 10.1-13.3 (12.2, average of 9), tarsus 14.7-17.3 
(QUE) Seated. 

Females (10 from Los Santos, Herrera, Veraguas, Province of Pan- 
ama, Darién), wing 41.0-46.5 (44.1), tail 35.3-43.8 (39.9), culmen 
from base 11.7-13.3 (12.2, average of 9), tarsus 15.3-16.7 (15.9) mm. 

Resident. Widespread and fairly common in a variety of mainland 
habitats including forest (although usually at breaks and edges), 
second-growth woodland, and scrub in semi-arid as well as humid areas. 
The Tropical Gnatcatcher is usually seen from sea level to about 1200 
m, the highest elevational record being a female collected on March 10, 
1951, by Frank Hartman at 1230 m on Cerro Pando, Chiriqui. Aldrich 
and Bole (Scient. Pub. Cleveland Mus. Nat. Hist., vol. 7, 1937, p. 23) 
found it one of the commoner birds in brushy scrub and forest mar- 
gins of the Tropical Zone, arid division (sea level to 300 m) on the 
western side of the Azuero Peninsula. Eisenmann (Smiths. Misc. 
Coll., vol. 117, no. 5, 1952, p. 48) calls it “not commen’ ‘oni Sarce 
Colorado Island in the Canal Zone. He has found it in many localities 
in the Canal Zone and in adjacent parts of the Province of Panama. 

The Smithsonian has specimens from nearly every section of the 
Republic, from Almirante, Bocas del Toro, and Puerto Armuelles, 
Chiriqui, in the west, to Jaqué in eastern Darién. P. p. superciliaris has 
been collected in Veraguas at Puerto Vidal, at the mouth of the Rio 
Vira, in the extreme southwest, and at Sona; in Herrera at Parita, 
Portobelillo, Santa Maria, and La Cabuya; in Los Santos at Las 
Palmitas and Pedasi; in Coclé at El Roble; in the Province of Panama 
from Bejuco in the west to Chiman in the east; and in Darien from 


FAMILY SYLVIIDAE ISI 


the mouth of the Rio Tuquesa on the Rio Chucunaque, as well as from 
Cana and Jaqué. There is a recent sight report of a pair on Rey in the 
Pearl Islands on February 11, 1970 (Ridgely, 1976, p. 279). Beyond 
Panama this race ranges north to Guatemala and probably south to 
adjacent northwestern Colombia. 

I found a fully grown young bird at Jaqué on April 13, 1946, so the 
nesting season in Panama may begin in Iebruary as it does in Costa 
Rica (Skutch, Pac. Coast Avif., no. 34, 1960, p. 34). On March 13, 
1948, I collected a pair in near breeding condition at La Cabuya, Her- 

‘rera. Along the Chepo Road in the Province of Panama, Major Gen- 
eral G. Ralph Meyer found a pair building a nest on March 18, 1944; 
the nest contained 1 egg on March 26, and 1 from the full clutch of 3 
eggs was collected on April 9. Bond and de Schauensee reported a male 
with testes enlarged that was collected at Garachiné, Darién, on April 
24,1941. On May 26, 1912, E. A. Goldman collected a female at Cana, 
Darien, with an egg almost ready to lay. 

Most of what is known of this race’s breeding biology comes from 
Skutch (op. cit. pp. 43-56). He found 6 nests, which ranged in height 
from 2 to 8 m off the ground. The nest is built by both male and female 
and closely resembles the lichen covered cup-shaped nest of the North 
American Blue-gray Gnatcatcher (P. caerulea). The clutch was either 
2 or 3, and the eggs white, finely speckled all over with brown. An 
egg obtained by Major General Meyer measured 14.4X11.7 mm. The 
males assisted in incubation, a habit shared with P. caerulea and P. 
melanura, and also with Ramphocaenus, but rare in tropical passerines 
as a whole. After a 13-day incubation period the young hatch entirely 
devoid of down. They are brooded regularly for at least the first 5 days 
and are fed entirely on insects. The length of the nestling period and 
the extent to which young are cared for thereafter are not known. 


POLIOPTILA PLUMBEA CINERICIA Wetmore 


Polioptila plumbea cinericia Wetmore, 1957, Smiths. Misc. Coll., 134(9), p. 80. 

(Isla Coiba, Panama.) 

Characters.—Similar to P. b. superciliaris, but dorsal surface, in- 
cluding the wings, decidely darker gray; lower foreneck and breast 
grayer (instead of white as in superciliaris); sides darker gray; bill 
averaging broader. 

Measurements.—Males (9 from Isla Coiba), wing 46.2-49.7 (48.5), 
tail 41.8-46.3 (44.1), culmen from base 13.0-14.9 (14.0), tarsus 16.6- 
yee (17.2) mm. 

Females (6 from Isla Coiba), wing 45.1-47.4 (46.6), tail 41.9-45.3 


182 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


(44.0), culmen from base 13.8-14.8 (14.4), tarsus 16.2-17.5 (17.0) 
mm. 

Resident. This race is endemic to Isla Coiba, where I found it to be 
common in January and February 1956, ranging in leafy cover from 
low second-growth thickets near the shore to the summits of the tallest 
forest trees in the interior of the island. Invariably they moved about 
among the twigs and leaves in unceasing activity in pursuit of tiny 1n- 
sects, often so high above the ground that I could barely detect their 
_ tiny forms. The slender body, with long, narrow tail held at an angle 
above the back, and their quick, nervous movements, mark them even 
when the gray and white plumage is not clearly seen. I found them in 
pairs, and near breeding at this season. On January 21 one male was 
much excited by my squeaking, and came to perch within 4 m of me 
while it sang repeatedly a series of high-pitched notes of the usual 
genatcatcher quality mingled with beautifully clear, warbling phrases of 
much louder sound that would have graced the gifted song of a mock- 
ingbird. 

Fisenmann (with E. S. Morton) on October 8-11, 1965, found this 
species more numerous on Isla Coiba than anywhere else in Panama. 
They also found it on nearby Isla Coibita. The vocalizations heard on 
Isla Coiba in October were neither musical nor noticeably nasal. Birds 
heard singing on Isla Coiba by Ridgely in early April 1976 seemed to 
him to vocalize like mainland birds. 


POLIOPTILA SCHISTACEIGULA Hartert: Slate-throated Gnatcatcher, 
Cazajején Gargantigris 


Polioptila schistaceigula Hartert, 1898, Bull. Brit. Orn. Club, 7, p. 30. (Cachabi 
[=Cachavi], 500 ft, Esmeraldas, Ecuador.) 


Small, slender; dark slaty gray, except lower belly to undertail co- 
verts, which are white. 

Description.—Length 97-102 mm. Adult (sexes alike), crown, back, 
and undersurface from throat to chest dark slaty gray; base of lores 
faintly white; upper throat lightly spotted white; broken eye-ring 
white; upper tail coverts slightly lighter gray; wings and tail black, 
outer two pairs of rectrices slightly tipped white; lower belly to under- 
tail coverts, underwing coverts, and inner web of underside of all but 
outer three primaries edged white. 

A male collected at the Peluca Hydrographic Station in Colon on 
February 27, 1961, had the iris reddish brown; maxilla and extreme 
tip of mandible black; rest of mandible neutral gray; tarsus and toes 
dark neutral gray; claws black. The tongue spots were tiny and incon- 


FAMILY SYLVIIDAE 183 


spicuous, being confined to the centers of the two posterior projections. 

Measurements.—Males (2 from Colén and Colombia), wing 45.0- 
45.8 (45.4), tail 40.7-41.5 (41.4), culmen from base 11.7, tarsus 15.2- 
17.5 (16.4) mm. 

Female (1 from Colombia), wing 45.5, tail 41.3, culmen from base 
10.5, tarsus 14.7 mm. 

Resident. Rare and little known. Hartert described this bird in 
1898 from a specimen collected in Ecuador. Wytsman (Gen. Av., Part 
17, 1911, p. 17) gives the range as northwest Ecuador (the type lo- 
cality, Cachabi) and Isthmus of Darién, possibly based on an unsexed 
adult in the Paris Museum collected by C. Viguier, as later reported by 
itelimeyvi (Cat. Birds Am. Field Mus. Nat. History, vols xiu, 1934, 
peo Griscom (Bull. Mus: Comp. Zool., vol. 78, 1935, p. 362) 
mentions only the Viguier specimen, and Eisenmann (Trans. Linn. 
BaemNman ol 7, 1955) p83) says (. Panama (Darien), Chapman 
(Bull. Am. Mus. Nat. Hist., vol. 55, 1926, p. 560), giving the range as 
northwestern Ecuador to northwestern Colombia, puts it in the Tropi- 
cal Zone. 

I encountered this bird once, at the Peluca Hydrographic Station on 
the Rio Boqueron in Colon, in March 1961. The bird worked in typical 
enatcatcher fashion through the tops of bushes on the bank of an open 
quebrada, moving with the usual vibrations of the tail and quick move- 
ments. Henry van Horn, who accompanied me on this trip, told me 
that he had seen 2 in similar situations traveling with groups of other 
small birds. The specimen I collected here anda male collected by Pedro 
Galindo at Cerro Quia (800 m) in southeastern Darién (Ridgely, 
1976, p. 279) are the only recent specimen records for Panama. 

Ridgely (im litt.) has seen this bird on two occasions in eastern 
Wesien 1) onl the/ slopes of Cerro Ouia (500)m) on’ July 17,1975; 1 
above Cana (670 m) on March 2, 1981. In both instances the birds 
were traveling with mixed flocks dominated by varions tanagers (Tan- 
gara, Heterospingus, Hemithraupis, etc.) in forest canopy, and were 
observed to forage mostly on terminal branches and in leafy areas, 
much as any other gnatcatcher. Neither bird vocalized. 


RAMPHOCAENUS MELANURUS RUFIVENTRIS (Bonaparte ): 
Long-billed Gnatwren, Cazajején Piquilargo 


Scolopacinus rufiventris Bonaparte, 1838, Proc. Zool. Soc. London, 5 (1837), p. 
119. (Guatemala. ) 


Small; bill long; upper surface brown; tail black, tipped white; 
throat white with dark gray streaks; rest of undersurface buffy. 


184 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Description.—Length 105-118 mm. Adult (sexes alike), crown 
and hindneck dark olive-brown; back and wings grayish olive; sec- 
ondaries and inner primaries edged cinnamon; tail graduated, blackish 
with all but central pair of feathers tipped white; facial area and sides 
of breast bright cinnamon; throat white with some dusky markings, 
varying in prominence; rest of undersurface buffy. 

Juvenile, like adult, but upper surface browner, undersurface pale 
grayish brown. . 

A male taken March 25, 1961, at Pacora, Province of Panama, had 
the iris light wood brown; maxilla and line on side of mandible fuscous- 
brown; rest of mandible pale brownish white; tarsus and toes dark 
neutral gray. 

Another bird taken at Canita, Province of Panama, had the iris dull 
brownish gray; maxilla fuscous, except base of culmen which is 
fuscous-black; rami brownish white; rest of mandible fuscous-brown; 
tarsus, toes, and claws neutral gray. 

Measurements.—Males (10 from Herrera, Province of Panama, 
Los Santos, San Blas, and Darién), wing 47.5-54.0 (50.0), tail 37.4- 
48.2 (41.7), culmen from base 22.7-25.7 (23.9), tarsus 19.3-21.1 
(20.1) mm. 

Females (10 from Herrera, Los Santos, Province of Panama, and 
Darién), wing 46.6-51.5 (48.5), tail 33.5-39.0 (37.3), culmen from 
base 21.6-24.8 (23.5), tarsus 19.7-21.8 (20.7) mm. 

Resident. Widespread and fairly common in the lower growth of 
forest edges, second-growth woodlands, and damp thickets of fairly 
humid lowlands through most of Panama. The range of rufiventris 
extends from southeastern Mexico to Colombia; other forms are found 
south through Brazil and Peru. In Panama, Ramphocaenus is usually 
found up to 1050 m (Ridgely, 1976, p. 280), but Frank Hartman col- 
lected a male at 1200 m on Cerro Pando, Chiriqui, on March 11, 1951 
(now in collection of Ohio State University). On the western slope of 
the Azuero Peninsula, Aldrich and Bole (Scient. Publ. Cleveland Mus. 
Nat. History, vol. 7, 1937, p. 25) found it uncommon in humid forest 
of the Tropical Zone (300-900 m). In the Canal Zone, Eisenmann 
(Smiths. Misc. Coll., vol. 117, no. 5, 1952, p. 48) found it “fairly com- 
mon in thickets in the lighter woods.” 

The Smithsonian has specimens from Bocas del Toro at Almirante; 
Chiriqui at Puerto Armuelles; Veraguas at Sona and Santa Fé; on 
the eastern side of the Azuero Peninsula at Punta Mala and Pedasi in 
Los Santos, and Pesé and El Barrero in Herrera; El Uracillo in Coclé; 
in the Province of Panama and Canal Zone at several localities from 


FAMILY SYLVIIDAE 185 


Bejuco in the west to Cerro Chucanti in the Serrania de Majé in the 
east; in Colon at Punta Pilon on the Rio Indio; San Blas at Mandinga, 
and Darién at Pucro and Jaqué. 

Ramphocaenus dwells in viny tangles of undergrowth and in lower 
trees where it moves wrenlike, cocking its tail, and flicking it constantly 
from side to side, rarely coming to the ground or foraging more than 
10 m above it. The diet, as evidenced by the stomach contents of a fe- 
male collected by Hallinan (Auk, 1924, p. 319) at Farfan, Canal Zone, 
is small insects and small seeds. Two collected by Burton (Bull. Brit. 
‘Orn. Club, 1975, p. 85) weighed 9.0 and 11.0 g. 

This bird is usually found singly or in pairs. Its song is a distinctive 
musical trill on a single pitch, sometimes rising and then lowering in 
volume at the end. A sharp wrenlike ticking and a low, dry churr are 
some of its other calls. 

In Panama the nesting season runs from at least late April to July. 
Willis, on Barro Colorado Island in 1961, saw a bird on its nest on 
April 23 and another building a nest on July 6 ( Willis and Eisenmann, 
Smiths. Cont. Zool., 1979, p. 25). Ridgely (in litt.) observed a pair 
feeding a very recently fledged young bird in the wind-swept elfin cloud 
forest on Cerro Jefe (1000 m), eastern Province of Panama, on July 
24, 1975. Bond and de Schauensee (Acad. Nat. Sci. Philadelphia, 
Mon. no. 6, 1944, p. 40) mention a female collected at Garachiné that 
had enlarged ovaries on May 8, 1941. A male I took at Sona, Veraguas, 
on May 25, 1953, was in full breeding condition. The only published 
description from Panama of the nest is of one found by Eisenmann 
(Auk, 1953, p. 369) in the Juan Franco suburb of the city of Panama 
on July 15, 1950, when it contained 2 almost naked young. The nest, 
an open cup, was in a damp thicket, near a stream, about 15 cm off the 
ground, ina small shrub shaded by trees. It was built among the shrub’s 
vertical shoots and was composed “chiefly of grass-stems, with a few 
twigs and dried leaves and to the exterior were attached several large 
dried leaves that hung loosely along the sides and extended below the 
nest proper, forming a sort of ornamental skirt.” The measurements 
were exterior diameter approximately 10 cm; interior diameter 8 cm; 
exterior depth 12.5 cm; interior depth 8 cm. 

Skutch (1960, Pac. Coast Avif., no. 34, pp. 54-61) found a nest of 
Ram phocaenus in Costa Rica on April 6, 1939, the first egg being laid 
on April 14 and the second the following day. These were “white, 
lightly sprinkled with fine, pale cinnamon spots over the whole surface, 
with these markings heaviest at the thick end.’’ Measurements were 
19.8X 14.3 and 19.1X13.5 mm. The incubation period at this nest was 


186 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


17 days and the young hatched without any natal down. Pinfeathers 
pushed through the skin when the nestlings were 2 days old, and at 7 or 
8 days they were all feathered. The young were fed small insects and 
spiders; they left the nest when 12 days old and able to fly only a few 
yards. 

I. O. Chapelle (1m litt. to Eisenmann) reports that in the Canal Zone 
he regularly found it with wandering bird bands (consisting chiefly of 
Iormicariidae) conspicuous because of its musical trill, which is ac- 
companied by tail vibration and sometimes by bill fluttering. The trill 
is usually preceded and sometimes terminated by a low chip. When 
foraging, the bird gives a drier, less musical version of the trill. 

Although Aldrich and Bole (Scient. Pub. Cleveland Mus. Nat. Hist., 
1937, vol. 7, 1937, p. 116) considered-the birds they collected in 
the western Azuero Peninsula to be almost identical to R. m. sanctae- 
marthae of Colombia, this race is actually paler and browner than are 
any Panamanian birds, all of which are referable to rufiventris. Speci- 
mens in the Smithsonian show the range of rufiventris to extend into 
Colombia in the departments of Choco, Antioquia, and Cordoba (So- 
corré, Rio Sint, and Quebrada Salvajin, Rio Esmeralda), whereas 
birds from farther north in Cordoba (Pueblo Nuevo, 14.4 km north 
of Planeta Rica; Nazaret, 19.2 km northwest of Tierra Alta) and east- 
ward through Bolivar, Magdalena, and Norte de Santander (Guama- 
lito, 7 km west of El Carmen) are referable to sanctaemarthae, the 
range of which is thus rather far removed from Panama, particularly 
the Azuero Peninsula. Some birds from the more westerly portions of 
Panama tend to have less rufous on the head than, for example, those 
from Darién, but these are darker and grayer on the dorsum than 
sanctaemarthae and should be referred to rufiventris. Van Tyne and 
Trautman (Occ. Papers Mus. Zool. Univ. Michigan, no. 439, 1941, 
p. 9) comment on the frequently remarked color variation in Middle 
American populations of this species. 


MICROBATES CINEREIVENTRIS (Sclater): Tawny-faced Gnatwren, 
Cazajején Carileonado 

Small, with long bill and short tail; upper surface brown; undersur- 
face gray with throat white. 

Description.—Length 94-102 mm. Adult (sexes alike), crown dark 
brown, back slightly lighter; wing coverts, primaries, and secondaries 
dark brown, edged rufous; tail dark brown, tipped blackish; side of 
face tawny; throat white, bordered by narrow black stripe on each side; 
upper breast striped black and white; rest of undersurface gray with 


FAMILY SYLVIIDAE 187 


central whitish streak more or less present; flank feathers olive-brown, 
fluffy, decomposed; undertail coverts whitish. 

This species, sometimes called the Half-collared Gnatwren, is found 
from Nicaragua to southern Peru. In Panama, where two subspecies 
occur, it is an uncommon and little seen denizen of thick undergrowth 
in humid forest in lowlands and hill country to perhaps 1000 m. Usually 
found in pairs or small family groups, its behavior is wrenlike. It hops 
actively from log to vine to twig, often clinging sideways, holding the 
tail almost at a right angle and habitually jerking or flicking it. Ordi- 
narily, it keeps low, not far above the undergrowth, but Ridgely re- 
ports seeing pairs foraging 5 to 7 m above the ground. 

The gnatwren’s diet is almost purely animal matter gleaned from 
plant stems and foliage or picked from the ground; sometimes it tem- 
porarily joins a mixed band of small birds following army ants. E. A. 
Goldman noted the stomach contents of 2 collected at Portobelo, Colon, 
in May 1911: both had full stomachs, one containing arachnid remains 
20%, bits of a bug 10%, ant remains 60%, moth remains 10%, the 
other with a spider 30%, 9 ants and 5 ant eggs 59%, another hy- 
menopteran 2%, bits of a bug 5%, elytra of a nitidulid 3%, embryo of 
emseediliZ/;. 

The gnatwren has a variety of calls suggesting a wren or an antbird, 
but nothing that resembles a territorial advertisement song. Ridgely 
(1976, p. 280) says the usual call is “a fast chattering chrichrichrich- 
richrv’ and Slud (Bull. Amer. Mus. Nat. Hist., vol. 128, 1964, p. 306) 
lists “a ‘woink, woink, woink,’ usually followed by a chatter,” “an 
antbird-like whining nasal ‘yeah,’ ” “an even ‘peep’ with the timbre of 
an antbird or flycatcher,” and several other sounds. Eisenmann re- 
ports that birds mist-netted deep within humid forest at 1000 m on 
Cerro Campana, aiter being caged gave a hissing ssssss. 

Olson (Proc. Biol. Soc. Wash., vol. 93, no. 1, 1980, pp. 68-74) has 
recently revised the taxonomy of this species. His review did not af- 
fect the nomenclature of any Panamanian populations. 


MICROBATES CINEREIVENTRIS SEMITORQUATUS (Lawrence) 


Ramphocaenus semitorquatus Lawrence, 1862, Ann. Lyc. Nat. Hist. New York, 
7, p. 469. (Line of Panama Railroad.) 


Characters.—Belly darker gray with little if any whitish suffusion in 
the middle; no postocular stripe. 

A male collected at the Peluca Hydrographic Station, Colon, on 
February 24, 1961, had the iris dark wood brown; mandible pale 


188 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


neutral gray; line of commisure on maxilla narrowly edged with 
neutral gray; rest of maxilla black; tarsus, toes, and claws dusky 
neutral gray. A male caught in a mist net at the head of the Rio Guabal, 
Coclé on February 26, 1962, had the iris dark brown; mandible light 
neutral gray; maxilla black; tarsus, toes, and claws dusky neutral gray. 
The tongue of one taken March 21, 1951, at Cerro Campana, Province 
of Panama, had dark spots on either side of the base like those of 
Ramphocaenus and Polio ptila but relatively smaller than in the latter. 

Measurements.—Males (10 from Province of Panama, Coclé, and 
Colon), wing 51.0-55.0 (53.0), tail 25.3-30.3 (28.4), culmen from 
base 17.4-19.4 (18.2), tarsus 21.1-25.8 (23.3) mm. 

Females (10 from Province of Panama and Colon), wing 48.8-55.7 
(51.8), tail 22.8-31.9 (26.7), culmen from base 16.9-19.9 (18.1), tarsus 
21.8-24.6 (22.9) mm. 

Resident. Uncommon in humid forest lowlands, occasionally up 
to 900 m, on the Caribbean slope from Bocas del Toro to San Blas, and 
on the Pacific slope at Cerro Campana and Cerro Azul. The Smith- 
sonian has specimens from Bocas del Toro: Changuena River (720 m); 
Coclé: head of the Rio Guabal at Tigre (475 m), and El Uracillo on the 
Rio Indio; Colon: Chilar, Boca del Rio Indio, Cerro Bruja (north side, 
600 m), and the Peluca Hydrographic Station on the Rio Boqueron; 
Province of Panama: Cerro Campana (south face, 900 m) and Cerro 
Azul; Canal Zone: Alajela; San Blas: Mandinga and Armila. 

Cory and Hellmayr (Cat. of Birds of the Americas, 1924, p. 211) 
give the range of this subspecies in Panama as “western Panama (Vol- 
can de Chiriqui, Santiago de Veraguas, Lion Hill Station).” Paynter 
(Check-list Birds World, vol. 10, 1964, p. 444) also says “western 
Panama,” without assigning birds of central Panama to any race. Ol- 
son (Proc. Biol. Soc. Wash. vol. 93, 1980, pp. 68-74) has found that 
this race extends along the entire Caribbean slope of Panama to north- 
western Colombia at Acandi on the Gulf of Uraba (USNM no. 
427204), and that Griscom’s (Bull. Mus. Comp. Zool., vol. 72, no. 9, 
1932, p. 366) identification of specimens from San Blas as the Colom- 
bian race magdalenae was erroneous. 

FE. A. Goldman found the nest and eggs of this bird on June 7, 1911, 
in Colon at Cerro Bruja, where it was “a rather common species at 
elevations up at least to 2000 feet.’’ His previously unpublished notes 
say “the nest was in the center of a loose mass of leaves and trashy 
material spread over an area one foot across on the flat, horizontal up- 
per side of a large palm leaf four feet from the ground. The nest was 


FAMILY SYLVIIDAE 189 


lined with plant fibers. Overhanging leaves above sheltered it from 
rain.” The nest contained 2 speckled eggs. 

The only other description of the nest and eggs is by Kiff (Condor, 
1977, p. 261-262), who found a pair carrying nesting material near 
mercer Viejo, I:leredia Province, Costa Ricaom April 3, 1971. Placed 
one-half meter off the ground in a broad-leafed shrub, the nest, when 
completed, was “an open cup with an exterior diameter of 10 cm. and 
exterior depth of 15 cm., the inner cup being 5 cm. in diameter and 4 

cm. deep. The outer walls were composed of green moss, papery bark 
_ fragments, leaf petioles, and bits of dead brown dicot leaves, all held 
together by plant fibers. The inner cup was lined with a soft layer of 
dead leaf skeletons, plant fibers, and a few strands of fine grass. The 
nest was attached by slender plant fibers to the trunk of the shrub 
along one entire vertical surface, and another side was built around a 
small limb which grew off the trunk at a sharp angle. The overall ap- 
pearance of the nest was that of a semi-suspended, bulky cup of green 
moss, partially concealed by the leaves of the shrub.” The eggs from 
this nest measured 18.8X13.8 and 18.5x14.2 mm. They were white 
“with a liberal sprinkling of fine reddish-brown and dark brown spots 
over their entire surface, but slightly denser on the larger ends.” Kiff 
considered the nest similar to that of Ramphocaenus and the eggs 
identical in coloration and shape. 


MICROBATES CINEREIVENTRIS CINEREIVENTRIS (Sclater) 


Ramphocaenus cinereiventris, Sclater, 1855, Proc. Zool. Soc. London, 23, p. 76, 
pl. 87. (“Pasto,” Colombia; Buenaventura substituted by Cory and Hellmayr, 
1924, p. 212.) 

Characters.—A distinct dark postocular stripe; underparts lighter; 
dorsum somewhat lighter, more olivaceous, contrasting with the darker 
crown. 

Measurements.—Males (10 from Darién and Colombia), wing 50.0- 
57.0 (53.0), tail 24.9-30.7 (27.8), culmen from base 17.5-18.8 (18.0), 
tarsus 19.5-24.4 (22.2) mm. 

Females (10 from Darién and Colombia), wing 49.5-54.5 (51.6), 
tail 26.7-30.4 (28.1), culmen from base 14.7-18.6 (17.2), tarsus 20.4- 
22:9 (21'6) mm. 

Resident. Uncommon in Pacific slope lowlands of Darién. De- 
spite the fact that Cory and Hellmayr (Cat. Birds Americas, 1924, p. 
212) give the range of this race as extending “from eastern Panama 
(Tacarcuna) along the Pacific coast of Colombia south to Chimbo, 


190 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Prov. Guayas, western Ecuador,” and Griscom (Bull. Mus. Comp. 
Zool., vol. 72, no. 9, 1932, p. 366) noted that nominate cinereiventris is 
“known to range northward to the Pacific slope of Darien,’ Paynter 
(1964, Check-list Birds World, 1964, p. 445) does not include Pan- 
ama in the range of this race. The Smithsonian has a series of 13 speci- 
mens from Jaqué and Rio Jaqué, at the mouth of the Rio Imamado, 2 
from Tacarcuna, and 2 from La Laguna that have the brown post- 
ocular stripe characteristic of nominate cinereiventris, the form resident 
on the Pacific slope of Colombia and Ecuador. 

Two specimens (Gorgas Memorial Laboratory Collection) were 
taken at Alturas de Nique, eastern Darién at “2000 ft.” Ridgely re- 
ports 2 seen in Cerro Quia cloud forest, 900 m, “with a very large 
mixed tanager flock . .. foraging 10-20 feet off the ground.” | 


Family MOTACILLIDAE: Pipits and Wagtails, 
Bisbitas y Lavanderas 


The pipits and wagtails are a worldwide family of 54 species, found 
primarily in the Old World. Two pipits inhabit temperate North 
America and two wagtails breed in Alaska but return to Asia each year — 
for the winter. Seven species of pipit including the Yellowish Pipit, 
represented by a race in Panama, occur in South America. Pipits are 
streaked, rather like larks, to match the grassy environment that most 
inhabit. The tail often is edged with white, making the birds suddenly 
conspicuous when flushed. Pipits walk rather than hop; the tail wag- 
ging habit is less pronounced in the Yellowish Pipit than in many 
others. Pipits feed mainly on insects, although some vegetable matter 
may be taken at times. Their song characteristically 1s given while in 
the air, often several hundred feet from the ground. Many species of 
temperate regions are migratory, and in Panama the Yellowish Pipit 
seems to some extent nomadic, since I have suddenly found groups of 
them in savannas that I had thoroughly searched for several days 
previously without success. 


ANTHUS LUTESCENS PARVUS Lawrence: Yellowish Pipit, 
Bisbita Amarillenta 


FIGurReE 16 
Anthus (Notiocorys) parvus Lawrence, 1865, Proc. Acad. Nat. Sci. Philadelphia, 
17, p. 106. (savanna near Panama.) . 
Rather small, upper surface brown streaked with buff; undersurface 
mostly yellowish white; outer tail feathers white. 


FAMILY MOTACILLIDAE IQI 


DescriptionLength 106-118 mm. Adult (sexes alike), crown, 
back, and wings blackish brown, with feathers edged yellowish or pale 
cinnamon-buff; rump and upper tail coverts tawny-olive; tail blackish 
brown, with outer two rectrices dirty brownish white, inner of these 
with brown edge to inner web, and next rectrix sometimes with whitish 
edge to outer web; lores and eye-ring pale buff; side of face speckled 
pale cinnamon and brown; throat yellowish white; breast streaked with 
dark brown and tawny-olive, sometimes extending on flanks; rest of 
undersurface yellowish white to buffy; underwing coverts buffy cin- 


~ namon and white. 


Juvenile similar, but upper parts spotted with pale brown; undersur- 
face brownish white, streaked on breast, narrowly on sides. 

A female taken January 22, 1963, at Penonomé, Coclé, had the iris 
brown; maxilla and tip of mandible fuscous-black; base of mandible 
brownish flesh color; tarsus and toes light buffy brown; claws dark 
mouse brown. 


Figure 16.—Yellowish Pipit, Bisbita Amarillenta, Anthus lutescens parvus. 


Measurements.—Males (10 from Chiriqui, Herrera, and Province 
of Panama), wing 58.0-61.1 (59.7), tail 41.3-45.6 (43.6), culmen from 
base 10.3-12.5 (11.6), tarsus 18.3-20.4 (19.3) mm. 

Females (8 from Chiriqui, Coclé, Herrera, and Province of Pan- 
ama), wing 56.7-60.5 (58.1), tail 40.8-44.3 (42.8), culmen from base 
10.2-11.9 (11.3), tarsus 19.2-20.8 (19.7) mm. 

Resident. This is a rather common though locally distributed bird 
in short grass savannas and fields in lowlands on the Pacific slope of 
western and central Panama. I have collected the bird at Las Lajas in 


192 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


Chiriqui, Santa Maria in Herrera, Penonomé and Anton in Coclé, and 
Chico, Pacora, and Chepo in the Province of Panama, the last-named 
locality being the easternmost at which this bird has been found. I 
have also seen it in Los Santos, at Pedasi. The westernmost record is 
of 2 males collected by W. W. Brown, Jr., on December 3, 1900, at 
Divala, Chiriqui. The race parvus is restricted to Panama. Other 
races of Anthus lutescens are found on the coast of Peru and Chile and 
in Savanna country east of the Andes from Colombia east and south 
locally to northern Argentina. 

The Yellowish Pipit is often the first bird visitors to Panama see, 
since it inhabits the landing fields of Tocumen Airport and Albrook 
Air Force Base; once when I landed at Albrook 2 of these pipits rose 
and flew along just beyond the wings of the plane, so close that I could 
see all their markings. Elsewhere, however, I have sometimes found 
them hard to detect—when strong winds blow across the short grass 
fields where they live, they prefer to crouch near the ground rather 
than fly. On the ground they walk and run rather than hop, and some- 
times wag their tail; their usual flight is deeply bounding, but I have 
also seen them rise 125 m in the air, circling in 300 m circles, occa- 
sionally setting the wings wide and sailing, after which they often 
closed the wings and dropped for several meters. When ready to come 
down they dropped directly to the earth or broke the distance in two or 
three sections. When in flight, they occasionally utter a low, double call 
like that of the temperate North American A. spinoletta, but somewhat 
harsher. The song, also given in flight is “initially a series of dzee’s on 
a rising pitch, usually given as the bird ascends in a looping manner 
into the sky, then a long slurred dzeeeeececeeee given as it slowly glides 
back to earth, often with a dzip at end.”’ (Ridgely, 1976, p. 280). Oc- 
casionally Eisenmann saw them also singing from the ground a shorter 
tsitsirrit and tsitzeeeeee. 

Yellowish Pipits nest in small colonies; in one at Santa Maria, Her- 
rera, I counted 15 or 20 birds. A pair with developing gonads collected 
at Penonomé, Coclé, on January 23, 1963, is the earliest indication I 
have of the onset of breeding. Others collected during February, 
March, and as late as April 20 were nearly in breeding condition, but 
on March 26, 1949, near Chico, Province of Panama I found a young 
bird just from the nest and on the same day collected adults that had 
not yet bred. On June 20, 1953, I took a female at Ancon, Canal Zone, 
that had recently laid, and on June 27 of that year I flushed a female 
from a nest at the La Jagua Hunting Club at Chico, Province of Pan- 
ama. The nest was on the ground, sheltered partly under a low grass 


FAMILY BOMBYCILLIDAE 193 


tuft, the top being covered. It was made of grasses and other soft 
vegetation, and contained 3 eggs. The female tried to decoy me with 
fluttering wings. The eggs were vinaceous-buff marked with wood 
brown; one had fine streaks all over the surface, slightly denser at the 
blunt end; the second had very fine spots and a ring of blotches at the 
blunt end; the third had fine speckles scattered at the pointed end, be- 
coming denser toward the blunt end. This last egg, the only one now 
unbroken, measured 17.3 x 14.0 mm. | 

J. R. Karr (Eisenmann im litt.) during a study in La Jagua grass- 
~ land (July 1968-July 1969), found three nests; September 12, 1968— 
2 eggs; January 4, 1969—2 young; January 15, 1969—3 eggs. 

Ridgely notes (im litt.) that this species appears to be quite par- 
ticular in its habitat requirements, and leaves areas that no longer suit 
it, only to return once again if they revert back. During the last 15 
years this cycle has taken place several times along the dike over the 
Tocumen marsh, which, when heavily grazed, provides good habitat, 
but otherwise does not and the birds are not present, even at the same 
times of the year. 

Two birds collected by Burton (Bull. Brit. Orn. Club, 1975, p. 85) 
at the military instruction center near Pacora weighed 13.0 and 13.5 g. 


Family BOMBYCILLIDAE: Waxwings, Chinitos 


The 3 species of waxwings breed in temperate Eurasia and North 
America. All are primarily fruit and berry eaters, also taking some 
insects on the wing; when not nesting they wander irregularly in flocks 
searching for food. The Cedar Waxwing is an irregular visitor to Pan- 
ama, not seen every year; it has been recorded as far south as Colombia 
and Venezuela. 


BOMBYCILLA CEDRORUM Vieillot: Cedar Waxwing, Chinito Cedroso 
FIGuRE 17 
Bombycilla cedrorum Vieillot, 1808, Ois. Amer., September (1807), 1, p. 88, pl. 57. 

(Amerique depuis le Canada jusqu’ au Mexique = eastern North America.) 

Medium size; slim, crested; mostly brown, with gray wings and tail; 
tail tipped bright yellow. 

Description.—Length 153-172 mm. Adult (sexes alike); lores 
black; black stripe through and below eye extending to rear of crown 
but not meeting; narrow white line surrounding black, including lower 
rear edge of eye; crown dark cinnamon, crested; upper back olive- 
brown; lower back, rump, and upper tail coverts gray; tail blackish 


194 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


gray, tipped lemon yellow; wing coverts olive-brown; primaries and 
secondaries blackish, with edge of outer web of all but outermost pri- 
mary edged gray and secondaries sometimes with waxy scarlet tips; 
upper throat black, fading to olive-brown, which continues to belly; 
belly pale greenish yellow; thighs gray; undertail coverts whitish; un- 
derwing coverts white. 

Immature, like adult but black only from maxilla to front of eye; 
crest and scarlet tips on secondaries usually lacking; undersurface 
broadly streaked olive-brown and creamy white. 

Measurements.—Males (10 from eastern North America, taken in 
May), wing 90.0-96.2 (92.7), tail 50.6-58.3 (54.5), culmen from base 
9.3-11.7 (10.4), tarsus 15.9-18.5 (17.1) mm 

Females (10 from eastern North America, taken in May), wing 
89.5-94.5 (92.6), tail 48.9-57.7 (54.3), culmen from base 9.5-10.7 
(10.1), tarsus 15.2-18.7 (17.0) mm. 

Migrant. A rare and irregular visitant, recorded some years from 
January to March from the western border east to the Canal Zone and 
the Pearl Islands. This species breeds in Canada and northern United 
States and winters irregularly south to the West Indies, and Middle 
America, casually to Colombia and Venezuela. Salvin and Godman 
(Biol. Centr.-Am., vol. 1, 1904, p. 216) said that at that time the south- 
ernmost occurrence of the species was in British Honduras, but Salva- 
dori and Festa (Bol. Mus. Zool. Anat. Comp. R. Univ. Torino, vol. 14, 
no. 339, 1899, p. 3) cite a specimen from Chiriqui collected by mirce: 
More recent Chiriqui records are those of Monniche (Blake, Fieldiana: 
Zool, vol. 36, no. 5, 1958, p. 552), who collected 5 at Quiel and Lérida 
(1560-1590 m) March 19-21, 1933, and of Frank Hartman, who col- 
lected a female February 13, 1953, at 1260 m on Volcan ‘de Chirigur: 
In the Canal Zone, Scholes and Scholes (Condor, 1954, p. 167) ob- 
served Cedar Waxwings five times at localities on the Pacific slope of 
central Panama in or near the Canal Zone from January 14 to March 
4, 1951, 1 near Farfan Beach, several near Chorrera, an unspecified 
number in Ancon, and a flock of about 15 near Camp Empire. My 
Own encounters with waxwings have been in the lowlands—on the 
Caribbean side at Almirante, Bocas del Toro, where on February 24, 
1958, I found a flock of 40 or 50 gathered in two tall trees, and on the 
Pacific side at Puerto Armuelles, Chiriqui, where on February 21 and 
26, 1966, I saw a dozen perched in a tall tree at the edge of a mangrove 
swamp. On San José Island in the Pearl Islands I saw a flock of 25 
flying north over the large open area near East Bay on February 29, 
1944. 


FAMILY BOMBYCILLIDAE I95 


More recently in the Canal Zone, Eisenmann (1m litt.) reports seeing 
two flocks, each with over 15 birds, on the Pipeline Road on March 7, 
1974, (with E. S. Morton) and several flocks aggregating at least 17 
birds, at Summit Gardens on February 6, 1973 (with Ridgely). J. J. 
Pujals reports it on Escobal Road on March 20, 1971. N. G. Smith 
reports seeing a group of 4 on the road above Santa Fé, Veraguas, 930 
m, on April 7, 1975, a late date. There seem to be no reports from the 
Azuero Peninsula or on the mainland east of the Canal Zone. 


“ 
SSH 
= SS 


4 
4 

TAL 

1 MP, 

WW) 

if! 

Y, G y 
ag y 

(PA) y t 


Figure 17.—Cedar Waxwing, Chinito Cedroso, Bombycilla cedrorum. 


F. G. Stiles and S. M. Smith (Brenezia, vol. 17, 1980, p. 149) be- 
lieve that occurrences in Costa Rica indicate it is irruptive at the south- 
ern end of its range, possibly with 3-4 year cycles—a major irruption 
occurred there in 1973-74, when birds were seen from December 1973 
to mid-May 1974 at various localities. 


196 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Family PLOCEIDAE: Weaver Finches, Pinzones Tejedores 


[PASSER DOMESTICUS (Linnaeus): House Sparrow, Gorrion Inglés 
Fringilla domestica Linnaeus, 1758, Syst. Nat., ed. 10, p. 183. (Sweden.) 


Up to about 20 House Sparrows were seen in downtown David, 
Chiriqui, in February and March of 1976 by C. Myers and Ridgely 
(in litt.); these appear to be established. Since at least March 1979 
small numbers have also been found breeding in Panama City, but as of 
1982 the population had declined (Ridgely, pers. comm.). On April 
29, 1980, Ridgely saw 3 at Changuinola, Boas del Toro. It is not cer- 
tain whether these birds are the result of local introductions or a spread 
from Costa Rica, where the population is slowly increasing. | 


Family STURNIDAE: Starlings, Estorninos 


[STURNUS VULGARIS Linnaeus: Common Starling, Estornino 
Sturnus vulgaris Linnaeus, 1758, Syst. Nat., ed. 10, vol. 1, p. 167. (Sweden.) 


A single individual was seen at Albrook Air Force Base, Canal Zone, 
on February 10-12, 1979, by J. and R. A. Rowlett (Ridgely, in litt.). 
This individual was probably released. | 


Family VIREONIDAE: Vireos, Peppershrikes, Shrike-Vireos, 
Vireos y Follajeros 


The Vireonidae is a strictly New World family that reaches its 
greatest diversity in the tropics but includes migratory forms from 
temperate regions of both North and South America. Although the 
peppershrikes (Cychlaris) and shrike-vireos (Vireolanius and Smarag- 
dolanius) were once placed in separate families (Cyclarhidae and 
Vireolaniidae), recent evidence from behavior and ecology (Barlow 
and James, Wilson Bull., 1975, p. 332) and anatomy (Raikow, Bull. 
Carnegie Mus. Nat. Hist. 1978, no. 7; Clark, Wilson’ Bullyigsiage: 
74) supports the opinion of Zimmer (Amer. Mus. Novit., no 1160, 
1942, p. 15) that these genera belong in the Vireonidae. Vireos are 
primarily insectivores that glean from foliage, branches, and trunks; 
some take fruit as well. Except for the small greenlets, Hylophilus, 
which often forage in an active, warblerlike manner, vireos move 
slowly and deliberately through the foliage, where their greenish plum- 
age makes them difficult to locate; their loud and persistent songs, 


FAMILY VIREONIDAE 197 


usually a monotonous warble or whistle, are given throughout the day 
and make these birds easy to find. All vireos, so far as known, build 
cup-shaped nests suspended in a forked branch in trees or shrubs. Six- 
teen species, including 1 peppershrike and 2 shrike-vireos, have been 
found in Panama. Six species in the genus Vireo occur in Panama as 
migrants or winterers from the north. The Yellow-green Vireo (Vireo 
flavoviridis) on the other hand is the only songbird that nests in Pan- 
ama and “winters” elsewhere. The other Panamanian species of vireos 
are permanent residents. 


KEY TO SPECIES OF VIREONIDAE 


1. Rufous stripe above eye. 
Rufous-browed Peppershrike, Cyclarhis gujanensis. p. 198 


INE EUIROCISESETIDECMOVEG EVE) 65% cha oc a cis cciuy celal tae eee eee eet ne cane Soave 7} 
ee UTUMOIMIE Ola CHOWAI ioe ein le hs ied aly ler aa Re Me ae en 3 
PSem MCAINERMeROMICEOwans | ELSA. 8s bin eo ca cen eet ane oh hee ales hate alae 4 


3. Yellow superciliary. 
Yellow-browed Shrike-Vireo, Smaragdolanius eximuus. p. 208 
No superciliary. 
Green Shrike-Vireo, Smaragdolanius pulchellus. p. 204 
4. Wing bars present 
NAT RI@? [ORES SDS evaltts Baten» MRR AR RANMA nee ee Pm CG dn ania es pee 7 
5. Throat bright yellow. 
Yellow-throated Vireo, Vireo flavifrons. p. 213 
AUTROAbaNAKbISite Of PAle wyiCllOW en ac NA se ee pees 5c bee slic aealua cosa ea #6 6 
6. Wing bars vellowish, partial eye-ring yellowish to white. 
Yellow-winged Vireo, Vireo carmioli. p. 211 
Wing bars white, supra-loral stripe white. 
White-eyed Vireo, Vireo griseus noveboracensis. p. 210 
PEESTIPETC MATA ESEME ea Skee Wold lc sl cba wie ukae A ae a oaks Uc hg 8 
SHO ME RCs yard SCI Da ue hot sare re cbs heute 4 gin een enca yee EIEN LSA LS or evlate a 12 
8. ‘Dusky moustachial stripe. 
Black-whiskered Vireo, Vireo altiloquus. p. 221 
RATA OUStaAGiNTal wb IDE ease vehi a. POS gM te cea MR ace ERS as i 9 
Ploincensuniace entirely, or partially ivellow....:c92eceus ele bah 6 ats cee 10 
Undersurface entirely white (sometimes pale yellowish wash on flanks). 
Red-eyed Vireo, Vireo olivaceus. p. 215 
10. Center of undersurface white, sides greenish yellow. 
Yellow-green Vireo, Vireo flavoviridis flavoviridis, p. 217 
Mo strotundersumiaceipal exvellowag ih use nein air Pee iain | 11 
11. Crown definitely brown, contrasting with more greenish back. 
Warbling Vireo, Vireo gilvus. p. 225 
Crown grayish olive, same color as back. 
Philadelphia Vireo, Vireo philadelphicus. p. 223 
12. Undersurface mostly white. 
Lesser Greenlet, Hylophilus decurtatus. p. 228 


198 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Undersuriace mostly:yellow or greenish. 1246). eee 13 
13. Forecrown definitely brown. 
Tawny-crowned Greenlet, Hylophilus ochraceiceps. p. 234 
Crown! greenish 00. 8 iso. se 6 paed on one ean ae a 14 
14. Lores yellow, crown mainly light olive-brown. 
Golden-fronted Greenlet, Hylophilus aurantifrons aurantitfrons. p. 231 
Lores and crown entirely olive-green. 
Scrub Greenlet, Hylophilus flavipes. p. 237 


CYCLARHIS GUJANENSIS (Gmelin): Rufous-browed Peppershrike, 
Pajaro Perico 


Ficure 18 


Tanagra gujanensis Gmelin, 1789, Syst. Nat., 1(2), p. 893. (French Guiana.) 


Large for a vireo; bill stout; head gray with rufous brow; rest of 
upper surface green; undersurface yellow and white. 

Description.—Length 134-148 mm. Adult (sexes alike), broad ru- 
fous stripe from forehead, above eye to rear of crown, thin line under 
eye rufous; crown and nape brownish gray, usually tinged with rusty, 
rest of upper surface olive-green; sides of head and hind neck gray; 
undersurface variable, with upper throat white or light neutral gray, 
rest of undersurface bright yellow becoming white below; underwing 
coverts and edge of wing yellow; inner webs of primaries edged with 
yellow. 

Immature similar, but superciliary stripe paler. 

The peppershrike is a variable species that ranges from eastern 
Mexico to central Argentina; 21 races are currently recognized (Blake, 
Checklist Birds World, 1968, pp. 103-107), of which 4 occur in the 
Republic. This is evidently an adaptable bird, since it dwells in quite 
different habitats in various parts of Panama, from forest borders in 
foothills and highlands of Chiriqui, at 840 to 2460 m, to light and scrub- 
by woodlands in lowlands on the Pacific slope in Veraguas and the 
eastern side of the Azuero Peninsula, and in mangroves on Isla Coiba. 
Although fairly common where it occurs, the peppershrike is always a 
hard bird to see, even when singing, as it perches among or moves 
slowly through the dense foliage, searching for its insect prey by eye 
rather than by the frequent hops of the smaller insectivores with which 
it sometimes travels. Were it not for its loud song, the peppershrike 
might be impossible to find altogether, as is evidenced by the vast pre- 
ponderance of males in museum series. The song is usually given re- 
peatedly, although I have also searched for individuals that paused for 
more than a half hour between songs. The song is a series of short, 


FAMILY VIREONIDAE I99 


musical, whistled phrases, each repeated for many minutes, until a new 
phrase is taken up and repeated, the entire performance lasting for 
hours. There is great variety in the total series of songs. 

Where it occurs in tall forest, the peppershrike often forages up 
high. In Costa Rica, Skutch (Publ. Nuttall Orn. Club, no. 7, 1967, p. 
124) several times observed peppershrikes feeding on fat caterpillars, 
which were held beneath one foot while pieces were torn off and swal- 
lowed, a habit shared with shrike-vireos and true vireos. The nest is 
likewise similar to that of the true vireos; the highest one Skutch found 
-was 9 m from the ground—although the birds foraged much higher in 
the surrounding trees—a cup hung by its rim from a forked branch, 
in typical vireo fashion. 


Figure 18.—Rufous-browed Peppershrike, Pajaro Perico, Cyclarhis gujanensis. 


The characters of each race found in Panama are given below, but it 
should be added that some forms intergrade; 2 birds from Aguadulce 
and FE] Potrero, Coclé, are intermediate between C. g. flavens and perry- 
got in the extent of yellow on the undersurface. 

On May 29-31, 1971, Eisenmann (im litt.) saw 2 or 3 individuals 
singing at Playa Coronado, western Province of Panama, in tall, 
planted trees, and a pair on visits on January 31, 1973, and March 27, 
1974. On earlier visits during the 1940’s to 1960’s when the natural 
vegetation was scrubby and planted trees were much younger and lower, 
he did not notice this species. The song resembled those heard in the 
western Chiriqui highlands. N. G. Smith told Eisenmann that he had 


200 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


observed the species in 1971 at Nueva Gorgona Beach, a few miles to 
the east of Playa Coronado. Another locality where the species has 
been found recently by Eisenmann and others 1s the hill country above 
Santa Fé, Veraguas. 


CYCLARHIS GUJANENSIS SUBFLAVESCENS Cabanis 


Cyclorhis subflavescens Cabanis, 1861, Journ. f. Ornith., 8 (1860), p. 405. (High- 
lands of Costa Rica.) 


Characters.—Gray of side of face continues behind crown to form 
band on upper back; chin and upper throat white; lower throat, upper 
breast, and flanks strontian yellow; lower breast to undertail coverts 
white. 7 

A male collected March 18, 1965, at El Volcan, Chiriqui had the iris 
light orange; maxilla light grayish brown, with distal end of culmen 
changing to dull black; mandibular rami dark neutral gray; anterior 
half of mandible ivory-white; tarsus and toes pale brown; claws light 
brownish neutral gray. 

Measurements.—Males (10 from Chiriqui), wing 71.8-76.8 (73.5), 
tail 54.4-60.1 (56.7), culmen from base 16.8- 19.8 (17.7), tarsus 20.9- 
A@ele( 2225) emai 

Females (5 from Chiriqui and Costa Rica), wing 72.4-77.5 (75.3), 
tail 56.7-60.1 (57.7), culmen from base 16.7-19.0 (17.8), tarsus 20.7- 
26.1 (22.4) mm. 

Resident. Fairly common in foothills and highlands of Chiriqui, 
840-2460 m. Known from the Volcan de Chiriqui ever since Arcé 
collected it there (Salvin, Proc. Zool. Soc. London, 1870, p. 184), it 
has also been collected at Boquete by W. W. Brown, Jr. (Bangs, Proc. 
New England Zool. Club, vol. 3, 1902, p. 59) and Davidson (Proc. 
Calif. Acad: Sei.; ser. 4, vol. 23, no. 17, 1938, p. 260)  aigiayewee 
countered peppershrikes at these localities and also at Cerro Pando. It 
is always highly vocal, but difficult to observe. 

In Costa: Rica, Skutch (Publ. Nuttall Orn: Club, nos ie7ae 
126-128) found nests between March 27 and early June. He describes 
the eggs as white, sparsely speckled on the thick end with small dots of 
bright brown. The only nest recorded from Panama was discovered 
by Worth (Auk, 1938, p. 539-540) on July 3, 1937, at El Volean (1230 
m). The nest was in a coffee grove in a large clearing in the jungle, 
and was built largely of a moss abundant on neighboring branches. It 
was attached to two forking twigs near the trunk, hung in typical vireo 
fashion, but woven more heavily and coarsely. The nest contained 2 


FAMILY VIREONIDAE 201 


fledglings about 5 days old, with pin feathers just beginning to sprout; 
evidently the young were born without down. They were fed entirely 
on insects, primarily soft caterpillars, and in the following week de- 
veloped rapidly, assuming the same plumage as their parents. Blake 
(Fieldiana: Zool., vol. 36, no. 5, 1958, p. 555) reported an immature 
bird with “the appearance of a recent nestling” collected by Monniche 
on the Volcan de Chiriqui on May 16. 


CYCLARHIS GUJANENSIS PERRYGOI Wetmore 


Cyclarhi. gujanensis perrygoi Wetmore, 1950, Proc. Biol Soc. Washington, 63, 
p. 61. (Ciénaga Macana, near El Rincon, Herrera, Panama.) 


Characters.—Lores light neutral gray; chin, extreme upper throat, 
and center of abdomen white; undertail coverts citron yellow; rest of 
undersurface strontian yellow, becoming lemon yellow on the lower 
breast and flanks. 

A male collected at Las Palmitas, Los Santos, on January 22, 1962, 
had the iris light orange; maxilla mouse brown; mandible neutral gray; 
tarsus, toes, and claws very light dull brown. 

Measurements.—Males (10 from Herrera and Los Santos), wing 
Vis 75,00 72.4),, tail 51.8-56.7 (54.2), culmen from base 16.6-18.9 
(17.5, average of 9), tarsus 20.2-23.3 (22.0) mm. 

Females (6 from Herrera and Los Santos), wing 65.8-76.2 (70.2), 
tail 51.6-56.6 (54.2), culmen from base 16.7-18.2 (17.1), tarsus 21.0- 
Za4 (21.9) mm. 

Resident. In light and scrubby woodlands in lowlands on the Pa- 
cific slope of western Panama from central and southern Coclé (Peno- 
nomé, Aguadulce) south through Herrera (El Rincon, Parita), and 
west in the drainage of the Rio Santa Maria into extreme east central 
Veraguas (F1 Villano, 24 km southeast of Santiago). Sight records 
assumed to be this race come from Santa Maria, Quebrada Tejel, and 
Quebrada Chitrabé to the south and west of Pesé, and FE] Barrero, 
ilertera, 

In the lowland, dry area of the eastern side of the Azuero Peninsula 
these birds are local in distribution, though not uncommon within the 
range outlined above. It is probable that they extend through tracts of 
scattered forest in hilly areas south through Los Santos, though there 
are no specimen records at present to substantiate this. On July 24, 
1964, Eisenmann and N. G. Smith saw 2 in Los Santos on a wooded 
stream border between Llano de Piedra and Tonosi; Smith saw 1 
carrying food. The brilliant coloration of this form is strikingly dif- 


202 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


ferent from that of C. g. subflavescens, found in Veraguas and Chiriqui 
to the west. The transition between these two forms in the lowlands 
east of Santiago, Veraguas, is apparently along the low watershed be- 
tween the Rio Santa Maria of Coclé, Herrera, and extreme eastern 
Veraguas, which flows into the Gulf of Panama, and the Rio Martin 
Grande, with its tributaries, which empties into the Gulf of Montijo. 
A specimen from E] Villano, in the drainage of the Santa Maria, is 
easily recognized as perrygoi, while another from La Colorada on a 
tributary of the Martin Grande, while somewhat intermediate, is to be 
placed with subflavescens. 

I have named this form for Watson M. Perrygo of the U. S. National 
Museum, my friend and companion on many days afield in Panama 
and elsewhere, who shot the first of our specimens of it. 


CYCLARHIS GUJANENSIS FLAVENS Wetmore 


Cyclarhis gujanensis flavens Wetmore, 1950, Proc. Biol. Soc. Washington, 63, p. 
174. (Chiman, Punta Madrofio, Panama, Panama.) 


Characters.—Gray of side of face continues behind crown to form 
band on upper back; chin and small area in center of abdomen white, 
rest of undersurface between lemon chrome and lemon yellow. 

Measurements.—Males (3 from Province of Panama), wing 69.2- 
73.8 (70.0), tail 49.0-52.3 (50.9), culmen from base 17.7-18:5) (18:1), 
tarsus) 20/5225:2) (2107) aaa 

Female (1 from Province of Panama), wing 66.0, tail 53.2, culmen 
from base 18.5, tarsus 21.6 mm. 

Resident. Known from coastal area of eastern Province of Pan- 
ama, with specimens from Chiman and Maje. 

In the course of fieldwork in eastern Panama in the dry season of 
1950 I was fortunate in securing specimens of the peppershrike, which 
on examination proved to represent an unrecognized form. The birds 
were found in small numbers in thickets back of the beach at the mouth 
of Rio Majeé, and at the mouth of the Rio Chiman, near Punta Madrono 
opposite Chiman; these localities are on the coast a short distance from 
the Darien border. It is probable that the birds I recorded in 1949 at 
Chepo on the lower Rio Mamoni are this race, and it is practically cer- 
tain that this is the form obtained in 1941 at Garachine by the Fifth 
George Vanderbilt Expedition (Bond and de Schauensee, Acad. Nat. 
Sci. Philadelphia, Mon. no. 6, 1944, p. 40). Likewise, a bird I found 
singing on Barro Colorado Island in the Canal Zone on March 9-10, 
1950—the only record to date for the Zone—and a pair Ridgely (1976, 


FAMILY VIREONIDAE 203 


p. 282) noted in mangroves near Tocumen, eastern Province of Pan- 
ama, on July 10, 1975, were probably representative of this race. More 
recently others have been seen at Juan Diaz; these are almost certainly 
flavens, spreading from farther east in the Province of Panama. 


CYCLARHIS GUJANENSIS COIBAE Hartert 


Cyclorhis coibae Hartert, 1901, Bull. Brit. Ornith. Club 12, p. 33. (Coiba Island, 
off Panama.) 


Characters.—Darker above than either subflavescens or perrygot; 
chin light neutral gray; upper throat to abdomen pale lemon yellow, 
brightest just above abdomen, then fading to white on abdomen and 
undertail coverts. 

A male collected on February 1, 1956, had the iris wax yellow; 
maxilla mouse brown; mandible neutral gray; tarsus and toes avil- 
laneous. 

Measurements.—Males (4 from Isla Coiba), wing 69.0-71.2 (70.6), 
tail 50.5-53.5 (52.6), culmen from base 16.4-17.6 (17.1), tarsus 21.0- 
22.4 (21.6) mm. 

I have found no female specimens in collections. 

Resident. During my stay on Coiba in January and February 1956 
the peppershrike was fairly common, but this was the beginning of the 
dry season and their songs had become infrequent; without these notes 
as a guide they are difficult to find. While they are robust in body, they 
move about behind leafy cover in such a leisurely manner, resting for 
minutes with only slight movements of the head, that it is only casually 
that one is seen. They are birds of the high forest crown, but come also 
about clearings, even into the low second growth called rastrojo, or to 
the borders of mangrove swamps. At the Maria work camp | found 1 
feeding in mango trees and coconut palms standing isolated in the ex- 
tensive clearing. 

The song is loud with strongly accented notes, and ends abruptly, 
when there is a pause of varying length, often of several minutes, be- 
fore it 1s repeated. The first two or three syllables are uttered rather 
slowly, followed by a rapidly given louder phrase. The notes carry for 
several hundred yards, and, if the song is continued, eventually the bird 
may be located, though the process of finding one may require half an 
hour. The 3 males that I collected represent many hours of search, 
since, as already stated, during January the birds were not singing 
steadily. 


204. BIRDS OF THE REPUBLIC OF PANAMA—PART ZA 


In the original description, written 80 years ago, when relatively few 
specimens were available, Hartert compared the Coiba bird with the 
race of Cozumel Island, off the coast of Quintana Roo, Mexico, which 
was suggested by the duller colors of cotbae. Actually, the subspecies 
of Coiba Island is more closely similar to the forms of the Panamanian 
mainland, from which it differs in very decidedly darker, duller colors, 
the breast and sides being distinctly greenish instead of bright yellow, 
the dorsal surface duller green, and the crown browner. The type speci- 
men (American Museum of Natural History), taken by Batty April 
20, 1901, an immature bird as is shown by the dark, almost black, bill, 
is browner on the crown than adult specimens. 


SMARAGDOLANIUS PULCHELLUS (Sclater and Salvin): 
Green Shrike-Vireo, Follajero Verde 


Ficure 19 


Vireolanius pulchellus P.L. Sclater and Salvin, 1859, Ibis, p. 12. (Guatemala. ) 


Large for a vireo; heavy-headed with stout, hooked bill; bright green 
with blue on hindneck and sometimes on head, and yellow throat. 

Description.—Length 126-144 mm. Adult (sexes alike), sides of 
head, face, and all of upper surface, brilliant parrot green, except blue 
hindneck and (depending on race) sometimes crown or forehead; 
abdomen and undertail coverts pale lemon yellow; edge of wing and 
underwing coverts, pale lemon yellow. 

The Green Shrike-Vireo is a common bird wherever it occurs in 
lowlands and foothills mainly on the Pacific slope, but it rarely descends 
from the tops of tall trees in forest and second-growth woodland, so it 
is usually not noticed by those who do not know its three- or four-note 
whistled song, which it delivers constantly. Occasionally it joins a 
mixed flock of insectivorous birds and descends to lower levels. Pan- 
ama is the southern limit of its range, which extends north to south- 
eastern Mexico. 

Lawrence (Ann. Lyc. Nat. Hist. New York, vol. 7, 1862, p. 468) 
received a specimen from McLeannan in 1862 and was the first to 
record this species from Panama, but not until 1905, when Ridgway 
described the race viridiceps from McLeannan’s specimen (Proc. Biol. 
Soc. Washington, vol. 16, 1903, p. 108), was it recognized that two 
forms exist in the Republic: one in the west with a blue forehead 
(verticalis), the other in the central and eastern region (viridiceps), 
with a green forehead matching the crown. The difficulty of collecting 


FAMILY VIREONIDAE 205 


or even seeing these birds at the many localities where they are com- 
mon makes it impossible even today to delimit the range of either race 
with any certainty. 


Figure 19—Green Shrike-Vereo, Follajero Verde, Smaragdolanius pulchellus. 


SMARAGDOLANIUS PULCHELLUS VERTICALIS (Ridgway) 


Vireolanius pulchellus verticalis Ridgway, Proc. U.S. Nat. Mus., vol. 8, 1885, p. 
24. (Angostura, Costa Rica.) 


Characters.—lorehead, hindneck, and sometimes area surrounding 
eye, light blue; upper surface parrot green. 

Measurements.—Males (6 from Veraguas, Costa Rica, and Nica- 
ragua), wing 68.0-76.5 (73.0), tail 44.4-52.4 (47.9), culmen from 
base 17.4-21.2 (19.0), tarsus 21.6-22.1 (21.8) mm. 

Females (4 from Veraguas, Costa Rica, and Nicaragua), wing 64.5- 
73.8 (69.5), tail 43.2-47.8 (46.1), culmen from base 17.0-19.5 (18.4), 
tansus 20.9-21.8 (21.3) mm. 

Resident. This race, which elsewhere occurs in Nicaragua and 
Caribbean Costa Rica, is known in Panama from 5 specimens. One is 
mentioned by Bangs (Proc. Biol. Soc. Washington, vol. 19, 1906, p. 


206 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


110), who says “I have one adult bird from Boquete, Chiriqui, that is 
absolutely typical.” This is probably MCZ no. 115194, sex ?, “Bo- 
queti,” Chiriqui, no date, from the Bangs collection. The catalogue 
gives no further detail, except an indication that the skin may have 
been entered about 1901. This bird has the forehead and band across 
the hindneck blue, and fully agrees with verticalis. 

Two others were collected at Calovévora, Veraguas, on the Carib- 
bean slope. A male collected by Arcé, now BMNH no. 85.3.10.208, 
has the blue between the lores typical of verticalis, as well as the nuchal 
stripe. AMNH no. 246930, collected by R. R. Benson, at Calovévora 
(marked Veraguas, but possibly extreme eastern Bocas del Toro) has 
the blue on the forehead present, but restricted in extent. AMNH no. 
187705-06, from Santa Fé, Veraguas, also collected by Benson, are 
typical of verticalis in the amount of blue on the forehead. These speci- 
mens are the basis for Griscom’s listing of the race in Veraguas (Bull. 
Mus. Comp. Zool., vol. 78, 1935, p. 363). 

A specimen (USNM no. 61990), collected by Arcé at Calobre, Vera- 
guas, has the feathers on the forehead partly missing, but the few that 
remain have a bluish cast in contrast to the bright green of the crown, 
thus showing an approach to verticalis. It appears that intergradation 
between verticalis and viridiceps takes place through the low divide of 
eastern Veraguas and probably western Cocle. 

The specimen cited in Salvin and Godman (Biol. Centr.-Amer., 
Aves, vol. 1, 1883, p. 210) collected by McLeannan at “line of railway” 
is probably the 1862 specimen Ridgway used as the type of S. /. 
viridice ps. 

N. G. Smith reports (in litt. to Eisenmann) hearing this species, 
presumably verticalis, in Bocas del Toro in forest near the Changuinola- 
Almirante Canal, during late March 1967. 

Slud (Bull. Am. Mus. Nat. Hist., vol. 128, 1964, p. 309) mentions 
various vocalizations and distinguishes between the songs of the two 
races, Saying verticalis has a musical usually three-note cheéur-cheéur- 
cheéur on the same level, while viridiceps has a three- or four- syllable 
chatter, chyuh-chyuh-chyuh-chyuh or an almost “spinking” pu-pu-pi. 
My own notes show that the song varies within Panama, but this con- 
stant singer is almost impossible to locate in the tall trees it inhabits and 
I have never been able to collect a singing bird. It is seen most often 
when foraging with a mixed flock. At Santa Clara, near Cerro Picacho, 
in Chiriqui I heard several giving a four-note song on March 16 and 
19, 1954. Near Puerto Armuelles, on the hillslope above the terminus 


FAMILY VIREONIDAE 207 


of the logging road at Corott the song I heard on February 12, 1966, 
was a three-note whistle. To the east, in the Canal Zone I have heard 
a three-syllabled whu-whu-whu at Juan Mina on January 10, 1955, 
and at Chiva Chiva on December 29, 1963. 


SMARAGDOLANIUS PULCHELLUS VIRIDICEPS (Ridgway) 


Vireolanius pulchellus viridiceps Ridgway, Proc. Biol. Soc. Washington, vol. 16, 
1903, p. 108. ( Panama.) 


Characters.—Blue only on hindneck; rest of upper surface parrot 
green, less yellowish green than in verticalis. 

A male collected at Charco del Toro, Rio Majé, eastern Province of 
Panama on March 27, 1950, had the iris brown; maxilla black; mandi- 
ble light neutral gray, darker toward base; tarsus, toes, and claws 
neutral gray; inside of mandible like exterior; tip of tongue horn color; 
rest of inside of mouth including basal part of tongue black; tongue 
deeply bifid. 

A female collected March 31, 1951, on Cerro Agua at El Valle, 
Coclé, had the dorsal pteryla broad in the center of the back and inter- 
rupted posteriorly so that the lower end presents a broad “V.” 

Measurements—Males (5 from Colon, Province of Panama, and 
Canal Zone), wing 67.8-71.2 (69.5), tail 43.0-48.1 (45.2), culmen from 
base 17.9-20.0 (18.7, average of 4), tarsus 21.1-23.6 (21.9) mm. 

Females (5 from Coclé; Province of Panama, and Canal Zone), 
wing 65.0-71.0 (69.8), tail 39.8-46.9 (43.7), culmen from base 16.8- 
20.5 (18.1), tarsus 20.4-22.2 (21.5) mm. 

Resident. Common in forest and tall second-growth woodland in 
the more humid areas on the Pacific slope from western Chiriqui to 
eastern Province of Panama; in the Canal Zone it has been commonly 
recorded from both slopes. It also occurs in southwestern (Pacific 
slope) Costa Rica. Specimen records are few, since this bird is ex- 
tremely difficult to collect as, when singing, it perches motionless in the 
tops of tall trees; fortunately it gives a distinctive three- or four-note 
whistle with what Chapman (My Tropical Air Castle, 1929, p. 240) 
calls “a tireless persistence.” The few specimens in the Smithsonian 
collections include 1 collected by Arcé at Calobre in Veraguas and re- 
cent ones from FE] Valle in Coclé, Charco del Toro, and Cerro Chucanti 
(at 510 m on the northwest slope of Serrania de Majé) in the Province 
of Panama; Cerro Galera, Albrook Air Force Base, and Barro Colo- 
rado Island in the Canal Zone; and the Peluca Hydrographic Station in 


208 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


Colon. A male collected at Puerto San Antonio, Rio Chepo (Griscom, 
Auk, 1933, p. 305) is in the Havemeyer Collection now at the Peabody 
Museum in New Haven. The American Museum has specimens from 
Santa Fé, Veraguas, and Cerro Azul, Province of Panama. 

I have encountered this shrike-vireo in Chiriqui at Puerto Armuelles, 
Buena Vista (750 m on the Rio Escarrea), and at Santa Clara, 1080 m, 
near Cerro Picacho; in Coclé at the head of the Rio Guabal (on the 
Caribbean slope) and at Fl Valle; in the Province of Panama at Canita, 
on Cerro Campana, and in Panama Viejo; in Colon on the Rio Bo- 
queron at 240 m and at the Peluca Hydrographic Station; and in many 
places in the Canal Zone including Chiva Chiva, Gamboa, Juan Mina, 
and Barro Colorado Island. 

Shrike-vireos stay in the densest leaves of the treetops, where their 
movements are so slow they are rarely noticed. It was not until March 
1950, when I was in the Serrania de Majé, where they were common, 
that I finally succeeded in collecting 1 and was able to associate with a 
particular species the song I had heard so often. Once, at Chiva Chiva, 
I watched a shrike-vireo foraging. Its green color matched the leaves 
exactly, with an occasional flash of blue from the neck. It searched the 
branches like a vireo, but moved far more deliberately, sometimes hang- 
ing underneath a branch while it examined the bark. Presently it seized 
a good-sized caterpillar, held it under its toes against the branch, and 
pulled it apart in sections, shaking its head to dislodge fluid from its bill. 

Although shrike-vireos sang constantly whenever I encountered 
them from late December through March, I found no evidence of breed- 
ing. Willis found a female building a mossy vireolike cup 14 m up as 
a male sang and watched, on May 15, 1966, in the Madden Forest Re- 
serve, Canal Zone ( Willis and Eisenmann, Smiths. Cont. Zool. no. 291, 
1979, p.25). This may be the first published information of the nesting 
of this species. 


SMARAGDOLANIUS EXIMIUS (Baird): Yellow-browed Shrike-Vireo, 
Follajero-Cejiamarillo 


Vireolanius eximius Baird, 1866, Rev. Amer. Birds, 1, p. 398. (“Bogota,” Co- 
lombia. ) 

Vireolanius eximius mutabilis Nelson, 1912, Smiths. Misc. Coll., 60, no. 3, p. 20. 
(Gana S000 tt easter) le amamiany) 


Large for a vireo; above bright green with light blue crown; super- 
ciliary and throat yellow; rest of undersurface light yellowish green. 


FAMILY VIREONIDAE 209 


Description.—Adult (sexes presumably alike), superciliary, spot 
at lower front of eye, and throat pale lemon yellow; lores blackish; 
crown light blue; rest of upper surface parrot green, slightly lighter 
on upper tail coverts; side of face and wing coverts green; primaries 
and secondaries black, with outer web edged green; tail green; under- 
tail coverts greenish yellow; rest of undersurface dull green-yellow; 
edge of wing, underwing coverts, and edge of inner web of primaries 
and secondaries pale lemon yellow. 

Measurements.—Type (female from Cana, Darién), wing 70.0, tail 
44.6, culmen from base 18.1, tarsus 22.1 mm. 

Resident. The only specimen known from Panama is a female 
collected at Cana (900 m), Darién, by E. A. Goldman on June 11, 1912. 
Nelson described this as the type of a new subspecies, mutabilis, that 
was said to differ from eximuius in having the supraloral part of the 
superciliary stripe broader, the forecrown greener, auriculars bordered 
with blue, yellow of throat changing abruptly into and contrasting with 
yellowish green of breast (however, there is a large patch of feathers 
missing from this area in the type), green of underparts paler, and 
undertail coverts brighter yellow. Blake (Check-list Birds World, 
vol. 14, 1968, p. 109) referred birds from Cordoba and Antioquia, 
Colombia, to mutabilis, but the series in the Smithsonian from this area 
is not separable from birds taken elsewhere in Colombia within the 
range of eximius. These specimens show that none of the characters of 
mutabilis are constant, the only possible distinguishing feature being 
the brighter undertail coverts of the type. Without additional speci- 
mens from Panama, it would be rash to conclude that mutabilis is a 
valid subspecies. It certainly does not constitute a “connecting link” 
with V. pulchellus viridiceps as postulated by Hellmayr (Cat. Bds. 
Americas, pt. 8, 1935, p. 189), and for this reason it seems prudent to 
maintain eximius as a distinct, monotypic species. 

Ridgely (im. litt.) has seen this species on two occasions in eastern 
Darién; 2 seen separately on slopes of Cerro Quia (530, 600 m) on 
July 17, 1975, and 1 seen above Cana (670 m) on March 2, 1981. All 
3 birds were foraging with mixed flocks in and just below the canopy. 
Ridgely never saw a bird sing, but believes that a shrike-vireolike song 
heard repeatedly must have been this species. It was distinctly more 
single-noted than is typical for pulchellus, e.g., pete-pete-pete. Of in- 
terest was his failure on either trip to even hear shrike-vireos at low- 
land locales; they were not present at Cana itself, only above. 

At Santa Fé, western Darién, on March 23 and 31, 1967, Eisenmann 


210 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


heard, coming from high forest trees, a song that he believed was ut- 
tered by Smaragdolanius. It was usually a three-noted pee-pee-pee, 
but occasionally two-noted. So far as he knows, no shrike-vireo has 
been collected or reported seen in this section of Darién. 


VIREO GRISEUS NOVEBORACENSIS (Gmelin): White-eyed Vireo, 
Vireo Ojiblanco 


Muscicapa noveboracensis Gmelin, 1789, Syst. Nat., 1(2), p. 947. (In Noveboraco 
[ = New York]; based on “Green Fly-catcher”’ of Pennant, 1785, Arctic Zool., 
2, p. 389.) 


Small; upper surface olive-green; yellow stripe from lores around 
eye; whitish wing-bars; undersurface white with yellow wash on sides. 

Description.—Length 112-124 mm. Adult (sexes alike), stripe from 
above lores and eye-ring yellow; upper surface plain greenish olive or 
olive-green; wings dusky, with middle and greater coverts tipped yel- 
lowish white, forming two wing bars; primaries and secondaries nar- 
rowly edged with greenish, and tertials broadly edged yellowish white; 
tail dusky with outer web of feathers edged greenish; throat and breast 
white lightly suffused with gray; sides and flanks pale lemon yellow, 
rest of undersurface white; underwing coverts and inner web of under- 
side of primaries and secondaries white. 

Measurements.—Males (10 from New Jersey, Delaware, and Mary- 
land, collected in May), wing 59.2-62.2 (61.2), tail 46.9-50.8 (48.5), 
culmen from base 9.9-11.9 (10.9), tarsus 17.5-19.7 (18.3) mm. 

Females (10 from New Jersey, Maryland, and Virginia, collected 
in May), wing 59.0-63.0 (60.6), tail 46.4-52.5 (48.5), culmen from 
base 10.7-12.8 (11.5), tarsus 15.9-19.8 (18.4) mm. 

Accidental. Known definitely in Panama from 2 specimens collected 
in Bocas del Toro. One, USNM no. 483154, a female, was caught on 
February 7, 1963, in a mist net in a fruit grove at Punta Vieja on Isla 
Bastimentos by C. O. Handley and F. M. Greenwell. They noted that 
the skull was ossified, the ovary measured 5X3 mm, and the iris was 
white. The second specimen, USNM no. 486555, unsexed, also came 
from a mist net, at Almirante, on October 16, 1964 (Hicks, Condor, 
1967, p. 90). This northern race of the White-eyed Vireo (breeding 
in eastern United States) normally winters south to Guatemala, north- 
ern Honduras, and western Cuba (A. O. U. Check-list of North Amer- 
ican Birds, 1957, p. 467); Bocas del Toro is more than 640 km south 
of its usual southern limit. 


FAMILY VIREONIDAE 211 


VIREO CARMIOLI Baird: Yellow-winged Vireo, Vireo Aliamarillo 


FicurReE 20 


Vireo carmiolt Baird, 1866, Rev. Amer. Birds, 1, p. 356. (Dota [ = Santa Maria 
de Dota], San José, Costa Rica.) 


Small; upper surface olive-green, with short yellowish eye stripe 
and broad yellowish wing bars; throat whitish, rest of undersurface 
yellowish. 

Description.—Length 109-116 mm. Adult (sexes alike), upper sur- 
face plain greenish olive; short, pale yellowish stripe from upper part 
of base of maxilla past eye, and yellowish border of lower rim of eye, 
forming a partial eye-ring; spot between lower part of base of maxilla 
and front of eye dark gray; sides of head yellowish olive; wing dusky 
with tips of middle and greater coverts tipped yellowish, forming two 
wing bars, outer web of primaries and secondaries edged yellowish 
Olive, and outer web of tertials edged yellowish white; tail black, 
feathers edged yellowish olive; throat whitish, tinged with olive yellow- 
ish; rest of undersurface light yellow tinged with olive on breast and 
sides; underwing coverts light yellow; inner edge of underside of pri- 
maries and secondaries white. 

Immature, like adult but head and nape dark cinnamon. 

A male taken February 24, 1965, at Volcan de Chiriqui, Chiriqui, 
had the iris reddish brown; maxilla and tip of mandible dull black; 
base of mandible dark neutral gray; tarsus, toes, and claws bluish 
neutral gray. 

Measurements.—Males (10 from Chiriqui and Costa Rica), wing 
64.0-67.5 (65.3), tail 44.7-49.6 (47.1), culmen from base 9.2-11.2 
(10.4), tarsus 16.9-18.6 (18.1) mm. 

Females (5 from Chiriqui and Costa Rica), wing 59.5-64.0 (62.0), 
tail 45.1-47.8 (46.6), culmen from base 9.9-10.5 (10.3, average of 4), 
tarsus 16.7-17.8 (17.3) mm. 

Resident. Fairly common in western Chiriqui in very humid forest, 
chiefly between 2000-2670 m, although recorded from 900 to 3170 m. 
Resident also in Costa Rica. W. W. Brown, Jr., collected a male at 
Boquete (1350 m) on March 17, 1901 (Bangs, Proc. New England 
Zool. Club, vol. 3, 1902, p. 59), but most of the localities from which 
this bird is known are higher—Blake (Fieldiana: Zool., vol. 36, no. 5, 
1958, p. 553), for example, reported them collected by Mo6nniche at six 
localities around Volcan de Chiriqui on the Boquete side between 1590 
and 2850 m. Two birds collected by Frank A. Hartman at Cerro 


212 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Copete (2100 m), above Boquete, are now at Ohio State University, 
and Davidson (Proc. Calif. Acad. Sci., ser. 4, vol. 23, 1938, p. 260) 
collected a male at Quiel (2340 m) above Boquete. I have collected 
Vireo carmioli on the west face of Volcan de Chiriqui at 2250 m, on 
Cerro Punta at 2100 m, and on’ Cerro Picacholat 2280 a: 


Ficure 20.—Yellow-winged Vireo, Vireo Aliamarillo, Vireo carmioli. 


Vireo carmioli is a bird of the high tree crowns, where I have some- 
times watched one move slowly in typical vireo fashion and then be- 
come more active, almost like a warbler; the large broad head and small 
bill give this bird a curious appearance. Eisenmann (Condor, 1962, p. 
505) reported only once seeing it as low as 13 m from the ground, but 
whether it normally nests that far from the ground is not yet known. 
On October 1, 1965, he saw 1 not 7 m up ina small roadside tree above 
Cerro Punta. The only nests that have been discovered are 2 found by 
Skutch (Publ. Nutt. Orn. Club, no. 7, 1967, p.\ 131) 1m@ostagisaes. 
both were in trees in pastures, 3 and 8 m from the ground. The diet 
seems to be insects and spiders gleaned from leaves and bark and, oc- 
casionally, picked from a leaf while the bird hovers before it. Some- 
times it joins mixed species flocks of other insectivores. 

These vireos seem to sing during much of the year; Skutch (of. cit.) 
has heard it in Costa Rica from early March to November. Ridgely 
(in litt.) has heard it in Panama in January. In Panama, Eisenmann 
(op. cit.) characterizes the song as a husky “cheéyah ...cheéyou... 
chipcheeweé, repeated a few times, with a long pause between repeti- 
tions.” Another song phrase or call is peeacheewit. 


FAMILY VIREONIDAE 213 


As mentioned above, the only information on this species’ nesting 
comes from Skutch’s observations in Costa Rica. The 2 nests he found 
were of typical vireo construction, a cup hung between two slender hori- 
zontal twigs, built of lichen, moss, bark, and leaf fragments, and silk 
from insect cocoons and spider webs. The first nest was begun on May 
21 (in Panama specimens collected by Monniche had enlarged gonads 
May 1-19 [Blake, op. cit.]) and the first egg was laid May 29. A second 
egg, which completed the clutch, was laid the next day. They were 
“white, with small, scattered dark spots on the broader end.” Both 
‘parents incubated, but the nest was destroyed before the eggs hatched. 
The other nest Skutch found contained young in late June; they were 
brooded frequently by both parents and were fed larvae. 


VIREO FLAVIFRONS Vieillot: Yellow-throated Vireo, 
Vireo Gargantiamarillo 


Vireo flavifrons Vieillot, 1808, Hist. Nat. Ois. Amer. Sept., 1(1807), p. 85, pl. 54. 

(United States [ = eastern United States].) 

Rather small; throat and breast bright yellow, rest of undersurface 
white; upper surface olive; yellow eye-ring; wings blackish with 
prominent white wing bars. 

Description.—Length 117-131 mm. Adult (sexes alike), supra-loral 
stripe, eye-ring, throat, and breast bright lemon yellow; side of head, 
and most of upper surface yellowish olive; wings blackish, with middle 
and greater coverts tipped white, forming two bars, undersurface below 
breast white. 

Measurements.—Males (10 from Panama), wing 70.3-78.5 (75.2), 
tail 44.2-50.4 (48.0), culmen from base 11.7-12.5 (12.2), tarsus 17.6- 
19.2 (18.6) mm. 

Females (10 from Panama), wing 72.0-77.2 (74.8), tail 46.0-50.1 
(48.3), culmen from base 11.1-13.2 (12.7, average of 9), tarsus 16.1- 
18.8 (17.6) mm. 

Winter visitor from temperate North America, known to winter 
south to Colombia. Widespread and fairly common, in every province 
of the Republic. I have also found them on several islands off the 
Pacific Coast, including San José in the Pearl Islands, Taboga, Parida, 
and Cébaco; | first thought these vireos were simply passage migrants 
on the islands, but on January 18, 1965, I found 1 ina berry tree on Isla 
Cébaco, which suggests by the date and source of food that they may 
winter there. As evidenced from the collecting localities, the Yellow- 
throated Vireo is found primarily in the lowlands, but in Chiriqui I 
have also collected it near El Volcan on the Silla de Cerro Pando at 


214 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


1500 m, and Blake (Fieldiana: Zool., vol. 36, no. 5, 1958, p. 553) men- 
tions a male collected April 4, 1933, at Lérida (1650 m). Griscom’s 
comments—“decidely uncommon winter visitant” (Bull. Mus. Comp. 
Zool., vol. 78, 1935, p. 364) and “few records south of western Pan- 
ama” (Amer. Mus. Nov. no. 282, 1927, p. 7)—do not seem accurate on 
the basis of more recent collecting and observation. Ridgely (1976, p. 
284) gives dates of occurrence from late October to early April. F. S. 
Blanton collected 2 males at Albrook Field in the Canal Zone on Oc- 
tober 22, 1953, and the latest spring occurrence of which I know is the 
April 4 bird from Chiriqui mentioned above. 

As a migrant and winter visitant in Panama the Yellow-throated 
Vireo’s habits are essentially the same as in their breeding areas to the 
north. They are slow moving even for a vireo, although they sometimes 
join groups of warblers moving through the trees. Willis (Living 
Bird, 1966, p. 205-206) observed 1 foraging near the ground on Barro 
Colorado Island on December 17, 1960, that ignored a nearby active 
swarm of army ants; he remarked that on Barro Colorado this vireo is 
usually found in the treetops. I have found them in mangrove swamps 
on San José, Pearl Islands, in open woodland along a quebrada in Ja- 
qué, Darién, and in shade trees over coffee at Santa Clara (1260 m), 
Chiriqui. At this season they take fruit as well as insects; 1 I shot on 
Isla Parida, Chiriqui, had the stomach filled with two good-sized drupes. 

Molt begins by January. A female I collected January 25, 1962, at 
Las Palmitas, Los Santos, was molting around the head; the incoming 
new feathers on the throat appeared very orange. A bird collected at 
the head of the Rio Guabal, in Coclé, on February 28 that year was in 
molt and had not developed body fat; another bird collected February 
28, 1965, at El Volcan, Chiriqui, also had no fat forming. Two birds 
that had just completed a molt and had no fat were collected at Santa 
Clara, Chiriqui, on March 17, 1954. I have collected specimens with 
heavy body fat on March 6, at El Potrero, Coclé, and March 17, at El 
Volcan. Ridgely (op. cit.) says its song is “Quite often heard, espe- 
cially in spring.” Eisenmann (in litt.) notes that this species gives a 
nasal snarl and a fast chittittittity, vocalizations also heard on the breed- 
ing grounds. 


[VIREO SOLITARIUS (Wilson): Solitary Vireo, Vireo Solitario 
Muscicapa solitaria Wilson, 1810, Amer. Ornith., 2, p. 143, pl. 17, fig. 6. (Bartram’s 
Woods, near Philadelphia, Pennsylvania. ) 
Vagrant. This species is included on the basis of three recent sight 
reports. At El Volcan in western Chiriqui, 1 was seen on March 17, 


FAMILY VIREONIDAE 215 


1972, by C. Leahy and R. Forster, another on January 16, 1974, by 
Leahy and Ridgely (Ridgely, 1976, p. 284). As the sightings were in 
exactly the same place they may have been of the same bird. Another 
bird was seen on January 24, 1977, near Fort San Lorenzo, Canal 
Zone, by R. Rodriguez (reported in Toucan, February 1977, p. 6, sup- 
plemented by a letter to Eisenmann). There are also recent sightings 
from Costa Rica. The race that normally occurs closest to Panama is 
nominate solitarius, which breeds in eastern North America and winters 
south in Central America in Nicaragua.| 


VIREO OLIVACEUS (Linnaeus): Red-eyed Vireo, Vireo Ojirrojo 


Rather small; upper surface olive- green with gray crown and white 
eye-stripe; lower surface white. 

Description.—Length 124-144 mm. Adult (sexes alike), crown gray 
bordered by narrow line of blackish; sides of face, rest of upper sur- 
face, including scapulars, and tail, grayish olive; green wing coverts, 
primaries, and secondaries dusky, with outer web of feathers edged 
yellowish olive; superciliary white, a narrow dusky gray stripe from 
bill through and behind eye; loral region below superciliary blackish; 
undersurface white; undertail coverts, sides, and flanks often tinged 
with yellowish olive; edge of wing white; underwing coverts very light 
yellow; edge of inner web of primaries and secondaries white on under- 
side. 

The Red-eyed Vireo of North America is a common migrant through- 
out Panama on its way to and from the Amazon Basin, where it spends 
the winter. It frequents clearings, second-growth woodland, and wood- 
land borders. Griscom (Bull. Mus. Comp. Zool., vol. 72, 1932, p. 366) 
wrote “I have yet to see any evidence to show that the Red-eyed Vireo 
winters with any regularity in any part of Central America,” and Eisen- 
miami ( onaiths: Misc. Coll, vol. 117, no.'5, 1952, p. 49)),"called it “a 
common migrant and occasional winter visitant in Panama, August 14- 
May 1, but chiefly seen in September and October, March and April.” 
Migration continues well into November. To date, there are no winter 
specimens. 

I. S. Morton (Eisenmann im litt.) saw a few on the Pacific slope of 
the Canal Zone in the final week of December 1970. Ridgely reports 1 
seen well on January 9, 1974, in forest on Barro Colorado Island. 

During both spring and fall the Red-eyed Vireo is “very abundant” 
at the banding station at Almirante (Loftin, Rogers, and Hicks, Bird- 
Banding, 1966, p. 41), often occurring in waves. In the spring of 1963, 
for example, Loftin and Olson (Carib. Journ. Sci., vol. 3, no. 4, 1963, 


216 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


p. 194) banded Red-eyed Vireos at Almirante between April 8 and May 
13. On most days no more than 6 were found in the nets, but on April 
18-19, 10 birds were taken each day and on April 28-30 from 10 to 18 
each day; after April 30 no more than 2 were netted each day. 

The large number of birds netted at Almirante has provided oppor- 
tunities to study the amount of energy expended by migrants. It is not 
known whether vireos reaching Panama in fall have flown nonstop 
from the United States or have come from farther north in Central 
America. Rogers and Odum (Wilson Bull., 1966, p. 421) found that 
newly-arrived Red-eyed Vireos had depleted their fat reserves and 
some were believed to have lost some of their fat-free weight as well. 
In an earlier study, however, Rogers (Bird-Banding, 1965, p. 116) 
collected a vireo killed in an October rainstorm at Almirante and esti- 
mated that on the basis of its remaining available fat it could still have 
flown another 658 km. A male I collected at La Jagua, Province of 
Panama, on March 20, 1961, was packed with fat. 

While in Panama, Red-eyed Vireos include fruit in their diet; from 
September to November 1968 Leck (Living Bird, 1971, p. 92) ob- 
served them repeatedly visiting a tree with mistletoe berries. In spring 
I have seen them moving through the trees with other migrant insecti- 
vores. 


VIREO OLIVACEUS OLIVACEUS (Linnaeus) 


Muscicapa olivacea Linnaeus, 1766, Syst. Nat., ed. 12, 1, p. 327; based mainly on 
“The red-Ey’d Fly-catcher, Muscicapa oculis rubris” of Catesby, 1731, Nat. Hist. 
Carolina, 1, p. 54, pl. 54, lower fig. (North America [| = (South) Carolina].) 


Characters.—Olive-green of upper surface darker than caniviridis, 
without grayish wash; pileum darker, mouse gray rather than olive- 
gray; undersurface with yellowish wash in sides and flanks. 

Measurements.—Males (10 from northwest Canada and United 
States, collected in May and June), wing 77.2-83.0 (80.6), tail 45.2- 
57.2 (54.4), culmen from base 12.3-14.7 (13.4), tarsus 17.1-19.1 
(17.8) mm. 

Females (10 from northwest Canada and United States, collected in 
May and June), wing 75.0-79.8 (77.1), tail 48.0-53.6 (51.1), culmen 
from base 13.0-14.5 (13.6), tarsus 16.4-18.8 (17.7) mm. 

Migrant and possibly winter visitant from the north. As a transient, 
Griscom (Bull., Mus. Comp. Zool., vol. 78, 1935, p. 364) called the 
Red-eyed Vireo common throughout. It has been recorded at sea level, 
on islands including Saboga, in the Pearl Islands (Thayer and Bangs, 
Bull. Mus. Comp. Zool., vol. 46, 1905, p. 154), and on Taboga (speci- 


FAMILY VIREONIDAE 217 


men in BMNH no. 1925.12.22.321), and at Almirante, Bocas del Toro, 
where it is one of the commonest migrants caught in the mist nets oper- 
ated by the Gorgas Memorial Laboratory. The highest elevation at 
which it has been collected is 1590 m, at Lérida, Chiriqui (Blake, 
Fieldiana: Zool., vol. 36, no. 5, 1958, p. 553). The Smithsonian has 
specimens from Jaqué, Darién; Chico, and La Jagua in the Province of 
Panama; from Alajuela and Fort Clayton in the Canal Zone; and 
from Almirante, Bocas del Toro. 


VIREO OLIVACEUS CANIVIRIDIS Burleigh 


Vireo olivaceus caniviridis Burleigh, 1960, Auk, vol. 77, p. 214. (Moscow, Latah 
County, Idaho.) 


Characters.—Olive-green of upper surface paler and with a gray 
wash; undersurface clearer white, with little or no yellowish olive on 
sides and flanks. 

Measurements.—Males (10 from Idaho, taken in May and June), 
wing 80.5-83.2 (81.8), tail 56.1-60.2 (58.0), culmen from base 14.1- 
16.4 (15.5), tarsus 17.1-20.0 (18.5) mm. 

Females (3 from Idaho, taken in July), wing 76.0-80.5 (78.3), tail 
56.2-58.0 (57.2), culmen from base 14.7-14.8 (14.7), tarsus 18.5-19.4 
(18.9) mm. 

Migrant from the north. Burleigh’s recognition of a separate race 
for Red-eyed Vireos from the arid northwestern United States, in 
Idaho, Washington, and Oregon, is not usually admitted (Blake, Check- 
list Birds World, 14, 1968, p. 122). Alan R. Phillips, however, con- 
siders this subspecies to be valid and has identified a male from the La 
Jagua Hunt Club, Chico, Province of Panama, taken March 30, 1955, a 
female from the mouth of the Rio Imamado, Rio Jaqué, Darién, taken 
March 30, 1947, anda female from Pacora, Province of Panama, taken 
October 14, 1958 as caniviridis. 


VIREO FLAVOVIRIDIS FLAVOVIRIDIS (Cassin): Yellow-green Vireo 
Vireo Verdiamarillo o Julian Chivi 
Vireosylvia flavoviridis Cassin, 1851, Proc. Acad. Nat. Sci. Philadelphia, 5, p. 152. 
(Panama and San Juan de Nicaragua; restricted to San Juan de Nicaragua by 
Zimmer, 1941, Amer. Mus. Novit., no. 1127, p. 2.) 
Vireo insulanus Bangs, 1902, Proc. New England Zool. Club, 3, p. 73. (Isla del 
Rey, Bay of Panama.) 


Rather small; crown medium gray; light gray superciliary; rest of 
upper surface bright olive-green; undersurface white, with flanks and 
undertail coverts yellow. 


218 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Description.—Adult (sexes alike), crown medium gray, sometimes 
with indistinct dusky margin; light gray superciliary; lores dull gray; 
rest of upper surface bright olive-green; sides of breast and flanks olive- 
yellow; undertail coverts bright sulphur or light lemon yellow; rest of 
undersurface white; edge of wing and underwing coverts bright sul- 
phur yellow. 

Juvenile, crown cinnamon-buff; whitish superciliary; rest of upper 
surface mainly olive-green; throat white; rest of undersurface very 
pale yellow. 

A breeding male collected March 22, 1961, at La Jagua, Province of 
Panama, had the iris light reddish brown; maxilla fuscous-brown; 
mandible dull ivory flesh color; tarsus and toes neutral gray. A female 
taken May 21, 1953, at Sona, Veraguas, had the iris light brownish 
orange; maxilla horn color; mandible light neutral gray; tarsus and 
toes dull neutral gray. The lighter iris seems definitely to be a sexual 
characteristic. 

Measurements.—Males (10 from Panama), wing 73.6-78.8 (76.7), 
tail 53.9-59.9 (56.7), culmen from base 14.2-16.3 (15.4), tarsus 16.9- 
NS C176) mina 

Females (10 from Panama), wing 69.0-75.9 (71.7), tail 46.5-53.1 
(49.5), culmen from base 13.0-15.2 (14.5), tarsus 17.1-18.6 (17.8) 
mm. 

Resident. Common to abundant in woodland borders, clearings, cut- 
over areas, coastal scrub and mangroves, gardens, open woods, and 
second growth on the Pacific slope and on islands off the coast and in 
extensively cleared areas of the Canal Zone to the Caribbean slope. The 
Yellow-green Vireo is a widespread breeder on the entire Pacific slope 
of the Republic, except in Darién, where its status is uncertain (pos- 
sibly only a transient). At higher elevations and humid lowlands it 
occurs only in cleared areas: W. W. Brown, Jr. (Bangs, Proc. New 
Iengland Zool. Club, vol. 3, 1902, p. 58) collected 4 at Boquete (1440 
m), and Blake (Fieldiana: Zool., vol. 36, no. 5, 1958, p. 553) records 
it from three localities around the Volcan de Chiriqui at 1560 to 1620 
m. In the western Azuero Peninsula, Aldrich and Bole (Scient. Publ. 
Cleveland Mus. Nat. Hist., vol. 7, 1937, p. 117) found it as high as 
900 m. Off the coast, the Yellow-green Vireo is very common on Isla 
Coiba and the Pearl Islands. On the Caribbean slope this species rarely 
appears away from cleared areas in the Canal Zone. Griscom (Bull. 
Mus. Comp. Zool., vol. 72, 1932, p. 366) notes a female taken by von 
Wedel on September 3, 1929, at Permé, San Blas, and Peters (Bull. 
Mus. Comp. Zool., vol. 71, 1931, p. 333) records another Wedel speci- 


FAMILY VIREONIDAE 219 


men taken September 27, 1927, at Changuinola in the Chiriqui Lagoon 
region of Bocas del Toro. Loftin (in litt.) reports that in 1962 Yellow- 
green Vireos were taken rather regularly between September 7 and 
October 30 in the mist nets operated by the Gorgas Memorial Labora- 
tory at Almirante, Bocas del Toro; presumably, these were migrants 
from farther north. 

The Yellow-green Vireo is the only oscine that breeds in Central 
America and is definitely known to migrate to South America, where it 
spends a few months in the upper Amazon Basin. Its breeding range 
extends north to Mexico and southern Texas; undoubtedly the birds 
mentioned above from the Caribbean slope and also some from higher 
elevations on the Pacific slope are transients. Yellow-green Vireos 
usually have passed through or withdrawn from Panama by late Oc- 
tober, although in 1963 Loftin netted 1 at Ancon, in the Canal Zone, as 
late as November 17. Although there is some variation from year to 
year, usually by late December (December 20, 1970, at Fort Kobbe, 
Canal Zone [Morton, Auk. 1977, p. 99]) the first birds return to the Pa- 
cific Coast; in mid-January birds returning there are at their peak. 
Farther inland the peak is slightly later, and on the Caribbean slope it 
does not occur until mid-March. (In 1962 I observed a considerable 
flight at El Potrero, Coclé, as late as March 8, when I saw as many as a 
dozen birds gathered in one berry-bearing tree at the same time). Mor- 
ton (op. cit) suggests that the breeding season is timed to coincide with 
an abundance of fruit, which is available sooner on the dry Pacific Coast 
than it is farther inland or on the Caribbean slope. The withdrawal to 
South America is at a time when fruit is more abundant there than 
during the late rainy season in Panama. 

H. Loftin’s banding data indicate that, despite migration, adults re- 
turn to the area where banded: 2 banded at Curundu, Pacific slope in 
the Canal Zone on March 6 and 9, 1963, were recaptured on February 
9 and 11, 1964, and the former again on April 2, 1966; 1 banded at San 
Francisco, Veraguas, on April 26, 1965, was recaptured May 5, 1969. 

The taxonomy of the Yellow-green Vireo is disputed. Blake (Check- 
list Birds World, vol. 14, 1968, p. 123) treats flavoviridis as a race of 
Vireo olivaceus, as well as the South American lV. chivi complex, al- 
though they differ in wing-formula and in color and there is no evidence 
of intergradation. Peters (Auk, 1931, p. 575-587) considered V. flavo- 
viridis as a full species, with the races insulanus breeding in Panama, 
nominate flavoviridis on the mainland north of Panama, and forreri 
breeding in the Tres Marias Islands off the west coast of Mexico, 
whereas Blake identifies some of the breeding birds of Panama as 


220 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


flavoviridis and included part of the Mexican mainland in the breeding 
range of forrert. When I was in the Pearl Islands in 1944 the bird was 
common and I secured a series of 21 skins to check on the question of 
the supposed form insulanus, described by Bangs (Proc. New Eng- 
land Zool. Club, vol. 3, 1902, p. 73) from El Rey Island and later con- 
sidered to be the bird of the Panama mainland also. There is variation 
in depth of color among these, some being decidedly darker, but this 
seems to be entirely individual, for which reason insulanus was synony- 
mized with flavoviridis (Wetmore, Smiths. Misc. Coll., vol. 106, no. 1, 
1946, p. 54). Migrant individuals of forreri may be recognized by 
their much larger size, as for example a specimen from Socorré, Rio 
Sinu, Cordoba, Colombia, taken by Carriker on April 25, 1949 
(USNM no. 412462). No specimens of this race have yet been iden- 
tified from Panama, however. 

Morton (op. cit., p. 101) examined the stomach contents of 98 
Yellow-green Vireos taken in Panama and found that 89 percent of the 
birds specialized in either fruit or insects, with their stomachs contain- 
ing 90 percent of one or the other; about half specialized in each, while 
the remaining 11 percent had stomachs containing equal portions of 
fruit and insects. Those specializing in insects had a far lower volume 
of food matter in their stomachs. The birds that filled up on fruit may 
be ones that learned the location of fruiting trees and visited them re- 
peatedly, even when the trees were outside of their breeding territory. 
The plants from which Morton found the vireos taking fruit included, 
in order of decreasing occurrence in stomachs: Miconia sp., Pitheco- 
lobium rufescens, Didymopanax morototom, Pittoniotus tricantha, 
Guazuma ulnufolia, Lasiacis sorghoidea. Two collected by Strauch 
(Bull. Brit. Orn. Club, 1977, p. 64) weighed 16.6 and 17.7 g. 

The arrival of the Yellow-green Vireo after its absence in South 
America is easy to detect, since it is a persistent singer, even through 
the hottest parts of the day. In tone and method of utterance the notes 
are like those of the Red-eyed Vireo, but somewhat more leisurely and 
slightly harsher in sound. Sometimes the phrases are shorter and the 
pauses in between longer than a Red-eyed Vireo’s, and I have also 
heard the song so harsh that it sounded almost like a House Sparrow’s. 
Morton (op. cit., p. 98) found that after pairing, males sang less per- 
sistently but gave rapid, songlike phrases when moving toward fe- 
males, who responded with a harsh chatter. In June and July singing 
increases somewhat, although often males were in molt and not associ- 
ating with females. They also give a harsh, nasal nywey (Eisenmann, 
in litt.). 


FAMILY VIREONIDAE 221 


During the courtship season I have several times observed males 
posed with head and neck erect, crest raised, and tail stiffly spread. 
Chasing is also frequent among males. Skutch (Pacific Coast Avif., 
no. 34, vol. 2, 1960, p. 22) describes a female Yellow-green Vireo tend- 
ing her nest while courted by an immature male that perched at the edge 
of the nest, swayed back and forth, and sang. In 1970 Morton found 
pairs on the Pacific slope of the Canal Zone beginning to nest in mid- 
January; they fledged young before any birds had even returned to 
the Caribbean coast, only 54 km away. On the Pacific Coast the last 
_ adults with dependent fledged young were seen on July 14. 

The nest is a typical vireo construction. I found one at La Palma, 
Los Santos, on March 27, 1948, that was hung in a little fork 5 m from 
the ground. The outside was covered with lichen and spider webs. 
Skutch (op. cit., p. 15) lists “small papery leaves of bamboo or grass, 
fibrous roots of epiphytes plucked from slender branches, and much 
cobweb” as the ingredients. The nest is built entirely by the female. 

The clutch is usually three eggs. Major General G. Ralph Meyer 
found a nest on May 15, 1941, on the Gamboa Road near the Naval 
Station in the Canal Zone; the measurements of all eggs were 21 X15 
mm. Peters (op. cit., p. 579) describes eggs of this species from Costa 
Rica as “white, speckled chiefly at the larger end with spots varying in 
color from a dark chestnut to an orange rufous, the chestnut pre- 
dominating.” 

Skutch (op. cit.) observed that incubation is performed entirely by 
the female and requires 13 or 14 days. At hatching the nestlings have 
a few scattered tufts of very short down and their eyes are tightly 
closed. They are fed a diet of insects by both parents. By their fourth 
day pinfeathers have begun to push through, but the birds’ eyes are 
still shut. On their eighth day of life the nestlings are feathered ex- 
cept on the head, and their eyes—brown rather than red—are fully 
open. At 11 days the young are well feathered and move restlessly in 
the nest, which they will leave within the next 3 days. 


VIREO ALTILOQUUS (Vieillot): Black-whiskered Vireo, Vireo Barbinegro 


Rather small; crown dull gray; rest of upper surface dull olive- 
green; superciliary whitish; dusky moustachial stripe; undersurface 
white; undertail coverts pale yellow. 

Description—Length 126-145 mm. Adult (sexes alike), crown 
brownish, usually without any dusky border; rest of upper surface 
dull olive-green; wing coverts, primaries, and secondaries dusky, with 


222, BIRDS OF THE REPUBLIC OF PANAMA—PART Al 


outer webs edged light yellowish olive; tail olive, usually with outer 
webs edged light yellowish olive; superciliary whitish to pale grayish 
buff; sides of face grayish buff; thin line above superciliary, line 
through eye, and moustachial stripe dusky; sides and flanks pale yel- 
lowish olive to grayish olive-green; undertail coverts pale yellow; rest 
of undersurface white; edge of wing very pale yellow; underwing co- 
verts yellowish white, edge of inner web of primaries and secondaries 
white on underside. 

The Black-whiskered Vireo breeds in southern Florida, the Bahamas, 
and the islands of the Caribbean. Two races are migratory, wintering 
in the Amazon Basin of South America. In Panama this vireo is known 
from 3 specimens and some recent sight reports. Ridgely (1976, p. 
285) refers to “several recent sightings in Canal Zone: One at Achiote 
Road on September 29, 1968 (Ridgely); and one at Gatun Dam on 
January 29, 1970 (E. S. Morton).” The specimen records are dis- 
cussed below. 


VIREO ALTILOQUUS BARBATULUS (Cabanis) 
Phyllomanes barbatulus Cabanis, 1855, Journ. f. Ornith., 3, p. 467. (Cuba.) 


Characters.—Superciliary stripe pale and less buffy than in /. a. 
altiloquus; throat and chest purer white. 

Measurements.—Males (10 from Florida and Cuba), wing 77.2- 
80.1 (78.5), tail 51.1-57.7 (55.8), culmen from base 14.7-17.1 (15.9), 
tarsus 15.1-19.5 (17.6) mm. 

Females (9 from Cuba), wing 72.0-78.2 (74.5), tail 49.2-55.3 (52.4), 
culmen from base 13.1-16.3 (15.2), tarsus 16.1-18.7 (17.5) mm. 

Accidental. This race breeds from the coast of southern Florida 
through the Florida Keys and Dry Tortugas to the Bahamas and Cuba, 
and winters in the Amazon Basin from northeastern Peru to north- 
central Brazil. Hasso von Wedel collected 1 (unsexed) at Puerto 
Obaldia, San Blas, on September 12, 1930, that Griscom (Bull. Mus. 
Comp. Zool., vol. 72, 1932, p. 366) considered “unquestionably this 
subspecies.” The specimen is now at the Museum of Comparative 
Zoology. Griscom (Bull. Mus. Comp. Zool., vol. 78, 1935, p. 364) 
lists this form from the Canal Zone as well, on the basis of a specimen 
sent to Salvin from there by McLeannan; at the time of Griscom’s 
writing that specimen had not been checked by him as to subspecies, 
but when I examined it at the British Museum in 1958 I found that, 
although in decidedly worn plumage, it agreed well with nominate 
altiloquus. 


FAMILY VIREONIDAE 223 


VIREO ALTILOQUUS ALTILOQUUS (Vieillot) 


Muscicapa altiloqua Vieillot, 1808, Hist. Nat. Ois. Amer., September, 1 (1807), 
p. 67, pl. 38. (Jamaica, Santo Domingo, etc.; types from St. Thomas, Virgin 
Islands, fide Stenhouse, 1930, Novit. Zool. 35, p. 271.) 


Characters —Back brighter olive-green than in V. a. barbatulus; 
superciliary less pale, more grayish buff; throat duller, chest tinged 
with pale grayish olive-green. 

Measurements.—Males (10 from Haiti), wing 73.2-87.5 (80.6), 
_ tail 48.7-57.5 (54.9), culmen from base 15.5-19.2 (17.0), tarsus 17.4- 
1967182) mm. 

Females (10 from Haiti), wing 74.2-81.0 (77.0), tail 44.5-55.5 
(51.2), culmen from base 14.9-18.9 (16.6), tarsus 17.0-19.8 (18.3) mm. 

Accidental. This race replaces V. a. barbatulus in most of the Greater 
Antilles and in the Virgin Islands; it winters mainly in northern South 
America, east to Guyana and south in Amazonia to lower Rio Madeira. 
Two specimens of this race have been collected in Panama. A male was 
sent by McLeannan to Salvin from the “Isthmus of Panama” (Sclater 
and Salvin, Proc. Zool. Soc. London, 1864, p. 348); it is now BMNH 
no. 85.3.10.1. Another male was collected by Benson on August 31, 
1926, at Guaval on the Rio Calovevora, on the Caribbean slope of Vera- 
guas and is now in the American Museum of Natural History (no. 


246926). 


VIREO PHILADELPHICUS (Cassin): Philadelphia Vireo, 
Vireo de Filadélfia 


Vireosyluvia philadelphica Cassin, 1851, Proc. Acad. Nat. Sci., Philadelphia, 5, p. 
153, pl. 10, fig. 2. (Bingham’s woods, near Philadelphia, Pennsylvania.) 


Small; gray crown, dull olive back; white superciliary; undersur- 
face mostly dull light yellow. 

Description.—Length 107-122 mm. Adult (sexes alike), crown plain 
mouse gray; rest of upper surface grayish olive-green, slightly lighter 
on rump; superciliary white; loral area below superciliary and spot be- 
hind eye dusky grayish; side of face pale olive; upper throat and lower 
abdomen whitish; rest of undersurface pale yellow, slightly deeper on 
breast; underwing coverts whitish. 

A female taken March 1, 1962, at the head of the Rio Guabal, Coclé, 
had the iris dark brown; maxilla and tip of mandible dark grayish 
brown; tip of mandible light neutral gray; base dull white; tarsus, 
toes, and claws dark neutral gray. 

Measurements.—Males (10 from Panama), wing 62.4-68.0 (66.3), 


224 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


tail 42.1-48.1 (45.9), culmen from base 10.1-11.9 (10.9), tarsus 12.8- 
16:7( 15:6); mami 

Females (10 from Panama), wing 61.0-69.2 (64.6), tail 38.1-45.5 
(43.2), culmen from base 9.9-11.7 (10.6), tarsus 13.3-16.9 (15.4) mm. 

Migrant from the north, and winter resident. Common in foothills 
and highlands of Chiriqui and Veraguas, rarer in lowlands of western 
and central Panama. The farthest east in Panama from which this 
species has been reported is in the Bayano River basin, on January 10, 
1976 (J. J. Pujals, to Eisenmann). The Philadelphia Vireo is found 
in Panama from October to mid or late April; usually it inhabits wood- 
lands, where it often travels with mixed flocks of North American 
migrants especially warblers, but I have also found it in mangrove 
swamps on islands off the Pacific Coast. The highest elevation at which 
it has been collected is 1620 m, at Lérida, Chiriqui, by Monniche (Blake, 
Iieldiana: Zool., vol. 36, no. 5, 1958, p. 554) and at Quiel, near Bo- 
quete, by Benson, and there are numerous records from elsewhere in 
the highlands of Chiriqui and Veraguas. I have collected Philadelphia 
Vireos in Chiriqui at elevations from 1260 m at El Volcan and Santa 
Clara, to Concepcion, 240 m, down to Las Lajas, Puerto Armuelles, 
and Punta Balsa, at or near sea level, and Isla Parida offshore. 

In the Veragus highlands, Arcé collected the Philadelphia Vireo at 
Chitra (Salvin and Godman, Biol. Centr.-Amer., Aves, vol. 1 (pt. 14), 
1881, p. 191) and Ridgely (in litt.) with F. Gary Stiles reports a “flocl< 
of at least 30 in woods along river below Santa Fé [1200 m.] in part as- 
sembled by Stiles’ pygmy-owl hooting” on January 6, 1974. My only 
encounters with the Philadelphia Vireo in Veraguas have, in contrast, 
been on islands off the Pacific Coast—-Cébaco, Gobernadora, Afuerita, 
and Coiba. 

On the western side of the Azuero Peninsula, Aldrich and Bole 
(Scient. Publ. Cleveland Mus. Nat. Hist., vol. 7, 1937, p. 118) found 
the Philadelphia Vireo “rather common in mixed flocks of arboreal 
North American migrants.’ Specimens at the Smithsonian from other 
provinces include birds from Almirante, Bocas del Toro; Santa Maria, 
Herrera; Pedasi, Los Santos; and Tigre, at the head of the Rio Guabal, 
Coclé. Ridgely (1976, p. 285-286) mentions some recent reports by 
Eugene S. Morton of the Philadelphia Vireo in the Canal Zone: “one 
collected at Albrook Air Force Base on February 19, 1971; one seen 
at Madden Lake Scout Camp on February 26, 1971; one collected at 
Gatun Dam on April 4, 1971.” 

Like many insectivorous migrants from North America, the Phila- 
delphia Vireo becomes more sociable out of the nesting season and more 


FAMILY VIREONIDAE 225 


flexible in its choice of habitat and food. It feeds extensively on in- 
sects, but at Isla Parida, Chiriqui, I took 2 on February 5, 1963, that 
had eaten small drupes. The specimen collected by Morton at Gatun 
Dam had in its stomach “red arils, green Micoma fruit; no insects,” 
while that taken at Albrook had “small beetles of several species and 
one small buprestid” (data on labels in American Museum of Natural 
History). On March 20, 1975, Morton and Eisenmann noted 2 in a 
Bursera tree with a variety of northern wood warblers in the light 
woodland of Farfan Road, southwestern Canal Zone. 

Buskirk et al. (Auk, 1972, p. 619), when studying mixed species 
flocks near Cerro Punta, Chiriqui found that Philadelphia Vireos were 
present in at least 26 percent of the flocks they observed; the vireos 
were “joiners,” but did not follow the flock from place to place. At 
the same locality Leck (Bird-Banding, 1975, p. 202) banded and 
weighed Philadelphia Vireos in 1968; he found that the average weight 
of 12 taken in March was 11.4 g and the next month 5 averaged 12 g. 
A bird that I took at Pedasi, Los Santos, on March 17, 1957, was not at 
all fat. 

Ridgely (1 litt.) observed a Philadelphia Vireo on Barro Colorado 
Island (March 13, 1979) with an antwren flock. He notes that in many 
areas this species seems to prefer coffee groves above all other habitats, 
especially where there are trees shading the coffee and where rank 
vegetation has grown up around the coffee bushes. 


VIREO GILVUS (Vieillot): Warbling Vireo, Vireo de Capucha Chocolate 
Muscicapa gilva Vieillot, 1808, Hist. Nat. Ois. Amer.; September 1, (1807), p. 65, 
pl. 34. (New York State.) 
Small; crown brown; rest of upper surface olive-green; superciliary 
white; throat and breast white; rest of undersurface pale yellow. 
Description—Length 107-121 mm. Adult (sexes alike), crown 
olive-brown; rest of upper surface grayish olive to brownish olive- 
green, superciliary white; dusky brown postocular streak and loral 
spot; facial area, throat, upper chest grayish, and white; rest of under- 
surface, edge of wing, and underwing coverts pale yellow or whitish. 
Vireo gilvus, the Warbling Vireo of North America and Mexico, in- 
cludes other forms through Central America and as far south as Bo- 
livia and southeastern Peru. Blake (Checklist Birds World, 1968, p. 
127-129) distinguishes 17 forms, all of which he puts in V. gilvus. 
Some consider the races south of Oaxaca, Mexico, distinct enough 
from northern gilvus to merit specific status under the name leuco phrys, 
while others believe the races breeding from southern Mexico to Nica- 


226 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


ragua are more closely allied to northern gilvus than to the forms from 
Costa Rica south. 

Little is known of the biology of any of the southern races. Slud 
(Bull. Amer. Mus. Nat. Hist., 1964, vol. 128, p. 312), writing of V. 
gilvus chiriquensis, the bird of Costa Rica and western Panama, says it 
generally occurs in small groups, in two’s, or singly, high in trees or 
lower in second growth and thickets. Like other vireos it 1s a persistent 
singer. Eisenmann (Condor, 1962, p. 505-506) found the song like 
that of a northern Warbling Vireo, but shorter and less varied, with 
song phrases lasting 3 to 3.5 seconds and pauses in between of 3 to 
5 seconds. Other vocalizations include a high thin tsip, sometimes 
dzheep; a tsip, tit-tit-tit-tit, given when moving about; and a sharp 
zweeyoo, zweeyoo, sweéyoo, given when gleaning, often hanging up- 
side down. 

Eisenmann (op. cit.) believes that V. gilvus has increased in Pan- 
ama with the clearing of forest. 


VIREO GILVUS CHIRIQUENSIS (Bangs) 
Vireosylva josephae chiriquensis Bangs, 1903, Proc. New England Zool. Club., 4, 

p. 9. (Boquete, 4000 ft, Volcan de Chiriqui, Panama.) 

Characters.—Back light grayish olive, contrasting with olive-brown 
crown; undersurface decidedly yellow. 

Measurements.—Males (10 from Chiriqui), wing 59.5-67.5 (64.0), 
tail 43.3-49.6 (45.9), culmen from base 10.4-12.6 (11.6), tarsus 15.3- 
1773) (165) mam: 

Females (10 from Chiriqui), wing 59.0-66.0 (62.0), tail 40.7-47.5 
(45.0), culmen from base 10.9-12.5 (11.9, average of 9), tarsus 15.4- 
1761720) roatan’ 

Resident. Common in forest borders, open woodland, and shrubby 
clearings in the highlands of western Panama, and in Costa Rica. 
Griscom (Bull. Mus. Comp. Zool., vol. 78, 1935, p. 364) gives its range 
in Panama as “mountains of Chiriqui and Veraguas.”’ Ridgely (1976, 
p. 286) includes western Bocas del Toro as well. W. W. Brown, Jr. 
(Bangs, Proc. New England Zool. Club, vol. 3, 1902, p. 58) collected 
this species at Boquete and on the Caribbean slope of Volcan de Chiri- 
qui between 1200 and 2100 m. Monniche (Blake, Fieldiana: Zool., vol. 
36, no. 5, 1958, p. 554) found it on Volcan de Chiriqui between 1560 
and 1860 m. Ridgely found it at 1150-1200 m at Fortuna, Chiriqui, in 
1976. Two specimens from Isla Cébaco and 1 from Isla Sevilla (both 
localities listed by Zimmer, Amer. Mus. Nat. Novit., 1127, 1941, p. 19), 
were collected by J. H. Batty, whose locality data has been shown to 


FAMILY VIREONIDAE 227 


be unreliable (Wetmore, Smiths. Misc. Coll., vol. 134, no. 9, 1957, p. 7). 
The specimens almost certainly come from Boquete, as did the re- 
mainder of Batty’s series in the American Museum. The Smithsonian 
specimens all come from Chiriqui—at Quiel (1560 m) near Boquete, 
Santa Clara (1260 m), Cerro Punta, and El Volcan (1350 m). 

My own experiences with this vireo have been limited. On March 
7, 1954, I shot 1 from a little group of migrant warblers in high 
branches at the border of forest near El Volcan. At other times I have 
found it in forest undergrowth. Eisenmann (Condor, 1962, p. 505) 
noted that at Cerro Punta it “favors borders of streams, edges of clear- 
ings, and cut-over open woods, rather than heavy forest.” In 1961 he 
found many singing in early April. My own observations were all in 
February and March of 1954 and 1955, when the notes I took made 
no mention of singing activity; I did remark, however, that a male 
collected March 22, 1954, was not breeding. The nest and eggs of the 
tropical forms of species have not been described. 

So far as known, the Warbling Vireo is insectivorous. Sometimes 
while foraging they hang upside down while among the terminal twigs. 
They participate regularly in interspecific foraging flocks. In addition 
to my own and Ejisenmann’s observations of this species in mixed 
flocks, Buskirk et al. (Auk, 1972, p. 619) found that at Cerro Punta 
Warbling Vireos were part of 25 percent of all flocks observed, al- 
though they never followed the wandering birds for any distance. 
Leck (Bird-Banding, 1975, p. 202), who weighed 8 birds that were 
caught in mist nets at Cerro Punta during September and October of 
1967 and another 8 the following March and April, found their average 
weight in “fall,” i.e., after the breeding season (12.8 ¢), higher than in 
“spring” during the breeding season (11.7 ¢). 


VIREO GILVUS DISSORS Zimmer 

Vireo gilvus dissors Zimmer, 1941, Amer. Mus. Novit., no. 1127, p. 18. (Cerro 

Munchique, 7000 ft, west of Popayan, Colombia. ) 

Characters.—Crown grayer, not warm brown of chiriquensis; upper 
surface greener, less brown; undersurface paler, less yellow. 

Measurements.—Males (10 from Colombia), wing 64.2-69.0 (65.9), 
tail 46.5-51.5 (48.6), culmen from base 10.9-13.9 (12.5), tarsus 16.2- 
17.3 (16.8) mm. 

Females (10 from Darién and Colombia), wing 62.0-67.9 (65.5), 
tail 45.6-51.3 (47.7), culmen from base 11.8-12.9 (12.3), tarsus 15.2- 
17.3 (16.4) mm. 


Resident. Known in Panama from a single specimen, a female col- 


228 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


lected on June 4, 1963, by Dr. Pedro Galindo at 1440 m, 6.4 km west 
of the summit of Cerro Mali, Darién. In Colombia this race—includ- 
ing Zimmer’s form disjunctus (Olson, Proc. Biol. Soc. Wash, vol. 94, 
1981, p. 363)—inhabits the entire extent of the western and central 
Andes. The specimen from Panama has the pale undersurface typical 
of Colombian birds and the upper surface most like that of dissors, to 
which it was tentatively assigned (Olson, ibid). 


HYLOPHILUS DECURTATUS (Bonaparte): Lesser Greenlet, 
Verdecillo Menor 

Very small; crown gray or yellowish olive; rest of upper surface 
yellowish olive-green; undersurface white, greenish yellow on flanks. 

Description.—Length 89-100 mm. Adult (sexes alike), crown either 
medium gray (slightly mixed with yellowish olive) or all yellowish 
olive; rest of upper surface yellowish green; wing coverts green; 
remiges dusky, with primaries edged narrowly and secondaries broadly 
green; tail green; narrow supraloral stripe and eye-ring whitish; side 
of face light gray; sides of breast and flanks olive-yellow; rest of under- 
surface white; edge of wing and underwing coverts pale yellow; under- 
side of inner web of primaries and secondaries edged yellow. 

Immature, undersurface tinged with buffy. 

The Lesser Greenlet is one of the commonest birds throughout the 
Republic, occasionally up to about 1800 m but is absent from drier 
scrubby or open areas on the Pacific lowlands of the eastern Azuero 
Peninsula, Coclé, and western Province of Panama. It frequents forest 
and second-growth woodland, where it gleans insects and spiders from 
middle and upper level foliage in the active manner of a warbler. It 
is a gregarious bird, often traveling in small groups, and a regular 
member of interspecific flocks of insectivores. At Jaqué, Darién, I 
once saw 1 with a little flock of the antwren Myrmotherula axillaris. 
Eisenmann (Condor, 1962, p. 507) considers it an ecological replace- 
ment of Vireo flavoviridis in the lowland canopied forest. He adds, 
however, that both occur together in lighter woodland and forest 
borders. 

The green-crowned populations of eastern Panama, Colombia, and 
western Ecuador, sometimes considered a distinct species, H. minor, 
intergrade in central Panama with gray-crowned H. decurtatus, of 
southern Mexico to western Panama. In the extensive series at the 
Smithsonian, most gray-crowned H. d. decurtatus show at least some 
green edging to the crown feathers. In the Province of Panama an ex- 
ample from Cerro Chame shows almost none, but series from Cerro 


FAMILY VIREONIDAE 229 


Campana and Rio Indio show slightly more than average. In an old 
series from Gatun, Canal Zone, all but one are gray-crowned with 
variable green edging to the feathers; the exception is all green and 
has been identified as H. d. darienensis. Two specimens from Juan 
Mina, Canal Zone, are nearly intermediate, but tending more toward 
gray. In 2 from Cerro Azul, to the east, 1 is an intermediate and the 
other darienensis. A specimen from Pacora still shows a trace of gray, 
while those from Chepo eastward are all darienensts. 

Ridgely (1976, p. 286) describes the call of this species as “a rapid 
musical phrase suggestive of a single phrase of the Yellow-green Vireo 
but even more monotonous, constantly repeated, typically deedereét or 
itsacheét.” Eisenmann (Smiths. Misc. Coll. vol. 117, no. 5, 1952, p. 
49) says the call is given at intervals of about 5 seconds. E. S. Morton 
(in litt. to Eisenmann) could find no constant difference between vo- 
calization in the Canal Zone of birds with predominantly gray crowns 
and those on Cerro Azul with completely olive-green or mixed crowns. 
A series he collected shows every variation between green and gray 
crown; moreover, seeming pairs included differently colored indi- 
viduals. 


HYLOPHILUS DECURTATUS DECURTATUS (Bonaparte) 


Sylvicola decurtata Bonaparte, 1838, Proc. Zool. Soc. London, 5(1837), p. 118. 
(Guatemala. ) 

Hylophilus pusillus Lawrence, 1862, Ann. Lyc. Nat. Hist. New York, 7, p. 323. 
(Atlantic side of Isthmus of Panama.) 


Characters.—Crown gray. 

A male taken February 18, 1966, at Puerto Armuelles, Chiriqui had 
the iris brown; maxilla except cutting edge fuscous-black; cutting edge 
and mandible pale neutral gray; gape dull honey yellow; tarsus, toes, 
and claws neutral gray; lower surface of toe pads dull honey yellow. 

Measurements.—Males (10 from western Panama), wing 50.8- 
94.4 (52.7), tail 32.6-38.9 (35.3), culmen from base 11.8-13.6 (12.7), 
tarsus 14.6-17.2 (16.3) mm. 

Females (8 from western Panama), wing 45.3-50.8 (47.7), tail 28.7- 
36.6 (32.5), culmen from base 11.7-13.8 (12.4), tarsus 15.2-16.8 (16.1) 
mm. 

Resident. Common in lowland forest, second-growth woodland, 
and coastal mangrove forest recorded on Volcan de Chiriqui as high 
esi S00) im) (Blake, Mieldiana:) Zool.) vol) 36))no. 5,’ 1958) pp. 554- 
955). Aldrich and Bole (Scient. Publ. Cleveland Mus. Nat. Hist., vol. 
7, 1937, p. 24) found it common in semi-deciduous coastal forest, be- 


230 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


low 300 m, near Montijo Bay, Veraguas. On Barro Colorado Island in 
the Canal Zone, where the population is intermediate between nominate 
decurtatus and darienensis, Willis and Eisenmann (Smiths. Cont. Zool. 
no. 291, 1979, p. 26) found it “very common in the treetops, often the 
center of mixed flocks there or in groups of its own kind.” The Smith- 
sonian has specimens from several locations in Chiriqui, including 
Puerto Armuelles on the coast and El Volcan at 1320 m; from Bocas 
del Toro at Almirante and Isla Bastimentos; Sona and Puerto Vidal in 
Veraguas; Pedasi in Los Santos; El Valle and El Copé in Coclé; and 
Rio Indio in Colon. 

There is little information on the breeding of this bird in Panama. 
E. A. Goldman’s notes include the mention of a female “in breeding 
condition” taken at Gattin, Canal Zone, on April 26, 1911. Stone (Proc. 
Acad. Nat. Sci. Philadelphia, vol. 70, 1918, p. 273) lists)” malewane 
young” collected at Gatun on June 25 and July 4, 1911. Fortunately, 
Skutch (Pacific Coast Avif., no. 34, 1960, pp. 29-34) has studied this 
race in Costa Rica. After living 20 years where Lesser Greenlets were 
abundant he found its nest in April of 1955. The nest was a deep cup 
attached by its rim to two diverging branchlets of a large muneco 
(Cordia) tree, 4.5 m above the ground. The structure was mainly 
leaves, in strips or fragments, or whole if small, all held together and 
attached to the branches by fibers and cobweb, and lined with a sparse 
layer of vegetable fibers. Outside dimensions were 8.9 cm in height by 
6.4 cm in diameter at the top; the interior was 4.5 cm in both diameter 
and depth. The eggs were not laid until a week after the nest was com- 
pleted. The two eggs, laid a day apart, were “white, scarcely glossy, 
spotted and blotched with pale brown, heavily on the large end and 
sparingly elsewhere. They measured 17.5 by 13.5 and 18.3 by 13.5 
millimeters.” 

Incubation took 16 days and when the young hatched they were de- 
void of down; the interior of their mouths was yellow. The young 
were periodically brooded by the female, but fed by both parents, who 
brought them insects and spiders. At 6 days of age the young had 
prominent pinfeathers, although still no down, and their eyes were 
partially open. Three days later feathers covered most of their bodies; 
at 12 days they left the nest, with 1 flying about 8 m in its first flight. 


HYLOPHILUS DECURTATUS DARIENENSIS (Griscom) 


Pachysylvia minor darienensis Griscom, 1927, Amer. Mus. Novit., no. 282, p. 7. 
(Cape Garachiné, eastern Panama.) 


Characters.—Crown yellowish olive. 


FAMILY VIREONIDAE 231 


A male collected March 9, 1963, at Armila, San Blas, had the iris 
mouse brown; maxilla fuscous-black; mandible neutral gray; tarsus, 
toes, and claws bluish neutral gray. 

M easurements.—Males (10 from Province of Panama), wing 50.1- 
fenn(a1/7)\, tail 3.2-33.7 (33.2), culmen from base 11-4-13.8 (12:8), 
tarsus 15.1-16.9 (15.6) mm. 

Females (10 from Province of Panama and Darién), wing 47.0-52.8 
(48.7), tail 26.1-35.4 (32.0), culmen from base 11.6-13.7 (12.6), tarsus 
15.1-16.9 (16.0) mm. 

Resident. Common and widespread in forest and second growth of 
eastern Panama, found in Darién as high as 1000 m on Mount Sapo 
and 1200 m on Cerro Pirre. This race is also found in Colombia south 
on the Pacific Coast to Dagua Valley, and the upper Sint and middle 
Magdalena Valleys. The Smithsonian collection includes specimens of 
“pure” darienensis from as far west as Utivé, on the Rio Colobre, in 
the Province of Panama, and also from Chepo, Chiman, and Cerro 
Chucanti farther east in that Province. The collection also includes 
birds from Armila, San Blas, and Jaqué, Cana, and Tacarcuna Village 
in Darien. 

Griscom (Amer. Mus. Novit. no. 282, 1927, pp. 7-8), when describ- 
ing this race wrote that in habits and song it is like decurtatus. Two 
stomachs from Cana examined by EF. A. Goldman held only insects: 
one had 3 ants 10%, caterpillar skin 10%, a large pale colored blattid 
80%; the other had bits of a hymenopteran 2%, a small moth with 4 
eggs 8%, a caterpillar 10%, bits of a small elaterid 5%, a small curcu- 
lionid 5%, a beetle 5%, other coleopteran remains 5%, 3 pale colored 
blattids (with 1 egg case) 60%. 

On February 22, 1950, I shot 3 Lesser Greenlets from a flock of 8 
or 9 at Chiman; of these, 2 were barely grown. Nothing, however, is 
known of the breeding biology of this race. 


HYLOPHILUS AURANTIIFRONS AURANTIIFRONS Lawrence: 
Golden-fronted Greenlet, Verdecillo Frontidorado 


Hylophilus aurantufrons Lawrence, 1862, Ann. Lyc. Nat. Hist. New York, 7 
(1861), p. 324. (Atlantic slope near Panama Railroad, hereby restricted to 
Frijoles, Canal Zone.) 


Very small; forehead brownish yellow fusing into brown crown; 
rest of upper surface yellowish olive-green; throat whitish; rest of 
undersurface light yellow. 

Description.—Length 98-107 mm. Adult (sexes alike), forehead 
yellow and olive-ocher, fusing into light olive-brown of crown; rest 


232 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


of upper surface, including wing coverts and tail yellowish olive-green; 
remiges dusky, with all but outermost primary edged yellowish olive- 
green, narrowly on primaries, broadly on secondaries; eye-ring white; 
sides of face light buffy brown; throat buffy whitish, faintly streaked 
with very pale yellow; faint tinge of light brownish buff across chest 
fused with pale yellow of rest of undersurface; sides and flanks pale 
olive greenish; underwing coverts and inner web of underside of 
remiges edged pale yellow. 

A. female collected January 12, 1963, at Chiva Chiva, Canal Zone, 
had the iris mouse brown; base of maxilla light fuscous-brown; tip of 
culmen dull black; cutting edge of maxilla and mandible flesh color; 
tarsus and toes neutral gray. 

Measurements.—Males (10 from Province of Panama and Canal 
Zone), wing 51.7-55.6 (54.1), tail 39.5-43.8 (42.0), culmen from base 
12.7-15.2 (13.8), tarsus 154-164) (15.8) mm: 

Females (7 from Province of Panama and Canal Zone), wing 48.6- 
53.5 (50.9), tail 34.5-42.6 (39.1), culmen from base 12.3-13.7 (13.1), 
tarsus 14.3-16.3 (15.4) mm. : 

Resident. Common on the Pacific slope, in Coclé (Aguadulce 
Salinas, Parita Bay), Herrera (southeast of Rincon) the Province of 
Panama (Playa Coronado, Nueva Gorgona, Cerro Campana, Panama 
City, Tocumen, Cerro Chame, Chico, Chepo) and in the Canal Zone 
(Rio Velasquez, Pedro Miguel, Corozal, Fort Clayton) east to south- 
western Darién (Garachiné), extending into the Caribbean drainage 
of the Rio Chagres to Culebra and Frijoles. It is also found on the 
Caribbean slope of northern Colombia; other races are found in eastern 
Colombia, Venezuela, and Trinidad. 

The single specimen from which this bird was named was sent to 
Lawrence by John R. Galbraith and James McLeannan during the 
“winter” of 1860-1861. Lawrence states that “the greater part of this 
collection was made on the Atlantic side of Isthmus, as the investigation 
of this section occupied their time until the season was too far advanced 
to enable them to procure many species on the western slope, and in the 
vicinity of Panama.” He then lists 6 species that were taken on the 
Pacific side. Since McLeannan is known to have collected in the vi- 
cinity of Frijoles, and since the bird is known to occur at that point 
through a specimen taken by Goldman, it seems appropriate to desig- 
nate this place as the type locality. From present knowledge this is the 
farthest point to which the bird penetrates in the Atlantic drainage. 

This small tropical vireo is found in open, thickety tree growth and 
scrubby vegetation, sometimes low down in the branches and sometimes 


FAMILY VIREONIDAE 232 


in the tops of the smaller trees, ranging from tree growth back of a 
mangrove swamp to the fringe of woodland bordering the course of 
streams, or the bases of hills rising in open savanna lands. I saw this 
species in shade trees in the clearings adjacent to the river, and Gold- 
man in 1911 recorded one in partly cleared jungle near Frijoles. The 
birds move about rather actively, suggesting small warblers in their 
movements, but usually remain behind the cover of leaves. Once I 
found one in company with a little group of the Lesser Greenlet. The 
song is not loud, but is interesting, usually a repetition of two notes, 
the second being stressed. Eisenmann (Condor, 1962, p. 507) describes 
the song as “distinctly vireonine, resembling that of H. decurtatus, but 
it is even less varied, most often with a four-syllabled whistled phrase, 
sliding down at the end—cheetsacheéyou or itsacheéyou—and with 
two-to-five second pauses between each repetition.” He reports also 
having heard a shorter witsachéw, and a more elaborate itsocheét cheet- 
sacheéyou, less musical than the vocalizations of H. decurtatus or of 
Vireo flavoviridis. This species also has a nasal scold, much like that of 
H. ochraceiceps, nyaah,nyaah,nyaah,nyaah, given in excitement. 

Eisenmann found the Golden-fronted Greenlet ecologically sym- 
patric with Vireo flavoviridis, although much more restricted in range 
and habitat, but tending to keep lower and avoiding the more humid 
second-growth areas. He adds that, as to other greenlets, in localities 
and habitats where he has found this species in Panama he has also 
found H. flavipes viridiflavus, and in localities with greater rainfall or 
better soil and taller tree growth or extensive woodland nearby (as in 
the Canal Zone and eastern Province of Panama) he has also found H. 
decurtatus, which avoids the drier areas and favors forest, just as H. 
aurantufrons avoids the more humid regions and favors scrub. E. S. 
Morton writes that in his experience, where the Golden-fronted Green- 
let and the Lesser Greenlet occur together, they often are found in the 
same group, and, although their songs, while of similar style, are 
readily separable, they have very similar or identical alarm and ago- 
nistic notes. 

On March 24, 1966, I watched 2 of these birds feeding on small 
berries at Chiva Chiva, Canal Zone. E. A. Goldman collected 1 at 
Corozal on June 15, 1911, whose stomach contained 2 or more ants 
60%, arachnid remains 30%, jassoid fragments 10%. Two collected 
by Strauch (Bull. Brit. Orn. Club, 1977, p. 64) weighed 9.6 and 10.1 g. 

Eisenmann found a bird building a nest in a small tree on a grassy 
slope with bushy vegetation while another sang nearby on June 17, 
1953, in the Juan Franco suburb of Panama City. 


234 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


HYLOPHILUS OCHRACEICEPS Sclater: Tawny-crowned Greenlet, 
Verdecillo Coronileonado 


Hylopilus ochraceiceps P.L. Sclater, 1859, Proc. Zool. Soc. London, 27, p. 375. 

(Playa Vicente, Oaxaca.) 

Very small; tawny brown; rest of upper surface brownish to olive; 
wings and tail russet; undersurface greenish yellow to olive-green. 

Description.—Length 95-112 mm. Adult (sexes alike), tawny 
brown sometimes with a bit of yellow at base of lores; rest of upper 
surface from dark russet-brown to olive-green; wing coverts as back; 
remiges dusky, with outer webs edged tawny brown; tail tawny brown; 
upper throat grayish white; rest of undersurface olive yellowish 
(bulunensis) or breast yellowish brown fading to light greenish yellow 
on remainder of undersurface; edge of wing, underwing coverts, and 
underside of inner webs of remiges yellow. 

The Tawny-crowned Greenlet is found from southeastern Mexico 
to northern Bolivia and Amazonian Brazil. Nine races are recognized, 
of which 3 occur in Panama; the Panamanian forms differ from one 
another mainly in the color of the back, which ranges from distinctly 
brown to olive-green. This greenlet is an uncommon resident of forest 
and second-growth woodland in lowlands and foothills; it has been 
recorded as high as 1500 m in Chiriqui (Ridgely, 1976, p. 287). Eisen- 
mann (Condor, 1962, p. 507) considers it more common in foothills 
than in the humid wooded lowlands. It inhabits dense undergrowth, 
rather than the higher levels in which the Lesser Greenlet moves; like 
other species of the genus, H. ochraceiceps is an active searcher for 
the small invertebrates that make up its diet. It often travels in pairs 
or family groups; it has been observed with various species of antwren 
and, in Belize, with Red-crowned Ant-Tanagers (Habia rubica) 
(Willis, Wilson Bull. 1960, pp. 104-105). Its voice is different from 
that of the other greenlets: Eisenmann (op. cit.) noted “a constantly 
uttered nya-nya, and a more vireonine weng,’ and Ridgely (op. cit.) 
mentions also “a rather long, slightly descending whistle.” 


HYLOPHILUS OCHRACEICEPS PALLIDPECTUS (Ridgway) 
Pachysylvia ochraceiceps pallidipectus Ridgway, 1903, Proc. Biol. Soc. Washing- 

ton, 16, p. 108. (Angostura, Cartago, Costa Rica.) 

Characters.—Upper surface definitely brown, breast with consider- 
able buffy tinge. 

A female taken March 15, 1965, at El Volcan, Chiriqui, had the iris 
light grayish brown; maxilla fuscous-brown; mandible dull neutral 
gray becoming browner on the distal end of the gonys. 


FAMILY VIREONIDAE 235 


Measurements.—Males (7 from Chiriqui and Costa Rica), wing 
52.0-59.0 (55.5), tail 40.8-44.8 (42.1), culmen from base 12.7-14.8 
(13.7), tarsus 15.2-17.4 (16.7, average of 6) mm. 

Females (6 from Chiriqui and Costa Rica), wing 50.3-57.0 (54.6), 
tail 39.9-46.0 (42.4), culmen from base 12.4-14.1 (13.4), tarsus 15.9- 
17.9 (16.9) mm. 

Resident. Griscom (Bull. Mus. Comp. Zool., vol. 78, 1935, p. 364), 
referring to this bird as nominate ochraceiceps, lists it for “Western 
Chiriqui (Pacific Slope).” Bangs (Proc. New England Zool. Club., 
vol. 3, 1902, p. 59) mentions a male collected by W. W. Brown, Jr., on 
the Caribbean slope of Volcan de Chiriqui, at 1200 m, on June 11, 1901. 
The only place I have encountered this race is at FE] Volcan, where on 
March 15, 1965, a pair flew into a mist net set at 1200 m at Palo Santo. 
Ridgely netted 2 (1 collected) on the upper Rio Chiriqui (Fortuna 
Dam site) at 1200 m on March 4, 1976. The only Bocas del Toro re- 
port is of 1 mist-netted by Loftin’s banders at Almirante on October 19, 
1965; the bird was examined in the hand by Eisenmann and released. 
Beyond Panama this race ranges to Honduras. 

Skutch (Pacific Coast Avif. no. 34, 1960, pp. 35-38) has watched 
1. o. pallidipectus at its nest in Costa Rica, where the breeding season 
starts at least as early as March. The 2 nests Skutch found were 2 and 
6.5 m off the ground; each was a typical vireo cup attached by cobwebs 
to a fork in a sapling. The nest he took apart for examination had an 
outer covering of moss, a layer of fine, light-colored bast fibers, a thick 
layer of long soft seed plumes, and a thin lining of more bast fibers. 
The 2 young in one nest observed by Skutch were brooded by the fe- 
male and were fed insects by both parents. The parents seemed to feed 
the young less frequently than do Lesser Greenlets, and the young left 
the nest at 13 or 14 days of age, slightly later than do young Lesser 
Greenlets. 

The only nest described from Panama was found by Eisenmann 
(Condor, 1962, p. 507) near El Hato, western Chiriqui on July 13, 
1949, but eggs had apparently not yet been laid; it was 1 m from the 
ground and covered on the outside by green moss. 


HYLOPHILUS OCHRACEICEPS NELSONI (Todd) 


Pachysylvia ochraceiceps nelsoni Todd, 1929, Proc. Biol. Soc. Washington, 42, 
p. 195. (Lion Hill, Panama). New name for P. o. brevipennis Nelson, 1911, pre- 
occupied by Helinat brevipennis Giraud, 1852 = Hylophilus decurtatus (Bona- 
parte, 1838.) 


Characters.—Upper surface brownish olive; less buffy below than 


236 BIRDS OF THE REPUBLIC OF PANAMA—PART 4A 


pallidipectus; more or less intermediate between that form and 
bulunensis. 

A male taken February 22, 1963, at Armila, San Blas, had the iris 
grayish white; maxilla black; mandible neutral gray; tarsus and toes 
bluish dark neutral gray. 

Measurements.—Males (7 from Veraguas, Province of Panama, 
Canal Zone, and San Blas), wing 54.5-57.0 (56.1), tail 37.9-45.8 
(40.5), culmen from base 13.5-14.5 (13.9), tarsus 15.1-16.5 (15.9) mm. 

Females (6 from Veraguas, Province of Panama, Canal Zone, and 
San Blas), wing 53.0-58.3 (55.6), tail 39.4-42.3 (40.7), culmen from 
base 12.2-15.2 (13.5), tarsus 15.1-17.0 (16.4) mm. 

Resident. Griscom (Bull. Mus. Comp. Zool., vol. 78, 1935, p. 365) 
lists nelsoni from the “Pacific slope, coastal forests of Veraguas to 
Rio Chepo, Darién [= Province of Panama]; rare.” I have collected 
this race beyond Griscom’s limits, at Cerro Chucanti (300 m), in the 
Serrania de Majé, eastern Province of Panama, and at Armila, San 
Blas. The Smithsonian also has specimens from Lion Hill and from 
between Summit Gardens and Gamboa in the Canal Zone, and from 
Chepo, Province of Panama. The Havemeyer collection, now in the 
Peabody Museum at New Haven, contains a male collected at Puerto 
San Antonio, on the Rio Chepo. 

EK. A. Goldman’s notes for May 1, 1911, when he was at Lion Hill, 
include the following: “A pair of these was seen and shot among low 
limbs of trees and rather thick jungle in fairly heavy forest. They were 
about 10 feet from the ground and were killed only about 15 feet apart. 
... They showed no breeding signs. Apparently a rare forest species.” 
The pair I found at Armila was caught in a mist net set in heavy forest, 
and 2 that I took at Cerro Chucanti were in tall tree tops. 

The American Museum collection contains a female (783985) col- 
lected January 23, 1964, by F. L. Chapman and E. Tyson in the Canal 
Zone 9.7 km north of Gamboa (Pipeline Road, Limbo Hunt Club) that 
contained an egg in the oviduct. 


HYLOPHILUS OCHRACEICEPS BULUNENSIS Hartert 

Hylophilus bulunensis Hartert, 1902, Novit. Zool., 9, p. 617. (Bulun, Esmeraldas, 

Ecuador. ) 

Characters.—Upper surface olive-green, undersurface olive yellow- 
ish, not yellow, with buff wash reduced. 

A male taken February 1961 at Cerro Pirre, Darién, had the iris 
brownish gray; maxilla and tip of mandible dusky neutral gray; rest 
of bill, tarsus, and toes neutral gray. 


FAMILY VIREONIDAE 237 


Measurements.—Males (7 from Darién, and Colombia), wing 53.0- 
61.8 (58.2), tail 38.0-46.3 (42.6), culmen from base 13.2-16.8 (14.5, 
average of 6), tarsus 15.5-16.7 (16.4) mm. 

Females (5 from Darién and Colombia), wing 50.5-55.9 (54.2), tail 
40.2-43.4 (41.5), culmen from base 12.7-13.9 (13.2), tarsus 15.1-16.3 
(i529). 

Resident. This race is known in Panama from extreme eastern 
Darién and San Blas. The Smithsonian has specimens from Darién 
taken at Jaqué, Cerro Pirre, Cana, and Tacarcuna Village. Bangs and 
Seagpeur Bull. Mus. Comp, Zool:, vol. 65, 1922, p. 223) mention '3 
collected at Mt. Sapo. Hasso von Wedel collected 2 females in San 
Blas at Obaldia and Ranchon that Griscom (Bull. Mus. Comp. Zool., 
vol. 72, 1932, p. 366) says “agree with three specimens from Mt. Sapo 
on the Pacific slope of Darién, but are perhaps a trifle less yellow below.” 
These locations are very close to Armila, where I collected a pair that 
are definitely referable to nelsoni, but as the Smithsonian also has speci- 
mens of bulunensis from the nearby Gulf of Uruba on the Caribbean 
coast of Colombia, it is possible that extreme eastern San Blas is an 
area where the two forms meet. In the Gorgas Memorial Laboratory 
collection are specimens taken on Cerro Quia, southeastern Darien, 
March 17, 1971, at 500 and 700 m. 

KE. A. Goldman examined the stomachs of 2 collected at Cana on 
June 6, 1912: 1 contained a locustid 25%, caterpillar remains (4 at 
least) 25%, 7 fragmentary ants 20%, 2 or more spiders including a 
ball of web 30%; the other contained a moth 8%, a lepidopterous co- 
coon 20%, ant remains 20%, arachnid remains 20%, a roach 32%. 


HYLOPHILUS FLAVIPES Lafresnaye: Scrub Greenlet, 
Verdecillo Rastrojero 


FIGURE 21 
Hylophilus flavipes Lafresnaye, 1845, Rev. Zool. [Paris], 8, p. 342. (““Bogota,” 

Colombia.) 

Small; entire upper surface yellowish olive; throat grayish; breast 
greenish yellow; rest of undersurface yellow. 

Description.—Length 105-117 mm. Adult (sexes alike), upper sur- 
face from crown to tail yellowish olive; wing coverts and secondaries 
yellowish olive; primaries blackish with outer webs edged yellowish 
olive; side of head light gray; chin dull white; throat and upper breast 
light yellowish olive, fading to light yellow on rest of undersurface. 

This species is found in the Tropical Zone from southwestern Costa 
Rica to the lower Rio Bayano, Province of Panama; it appears again 


238 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


in northern Colombia and ranges across Venezuela from the Orinoco 
Valley to the north coast, including Margarita Island, and the island of 
Tobago. The subspecies found in Panama are separated widely from 
those of Colombia and Venezuela by the forest of Darién and the Atrato 
Basin. 

The bird of Panama is much brighter colored above and below than 
its more eastern (South American) relatives and in fact is so different 
that the decision to place it as a geographic race instead of a distinct 
species is more or less an arbitrary one, based on general similarity and 
its allopatric range. 


YE; B = 
LZ ZZE 
GEE 
7 Z 
WN z= 
TY = eG 


Ficure 21.—Scrub Greenlet, Verdecillo Rastrojero, Hylophilus flavipes. 


HYLOPHILUS FLAVIPES VIRIDIFLAVUS Lawrence 


Hylophilus viridiflavus Lawrence, 1862, Ann. Lyc. Nat. Hist., New York, 7 (1861), 
p. 324. (Atlantic slope near Panama Railroad, Canal Zone, Panama, hereby 
restricted to Frijoles, Canal Zone.) 

Hylophilus viridiflavus pallescens Davidson, 1932, Proc. Washington Biol. Soc., 
vol. 45, p. 168. (Concepcion (1500 ft), Chiriqui, Panama.) 


Characters.—Undersur face light yellow. 

A male collected January 15, 1955, at Juan Mina, Canal Zone, had 
the iris cadmium yellow, bill light hair brown; line of culmen fuscous; 
tarsus, toes, and claws light yellowish brown. A male collected March 
24, 1961, at Chico, Panama, had the iris pale yellowish white; maxilla 
except cutting edge mouse brown; mandible and cutting edge of maxilla 


FAMILY VIREONIDAE 239 


pale brownish white (‘flesh color”); tarsus, toes, and claws light 
mouse brown. Another male, taken March 7, 1962, at Il Potrero, 
Coclé, had the iris light buffy gray; line of culmen and side of mandible 
to rami dull brownish white; gonys and underside of mandible light 
brownish white; gape dull honey yellow; tarsus, toes, and claws pale 
brown. 

Measurements—Males (15 from Panama), wing 54.5-58.3 (56.0), 
tail 44.8-49.8 (47.9), culmen from base 13.1-14.2 (13.6), tarsus 17.6- 
19.6 (18.7) mm. 

- Females (11 from Panama), wing 52.0-54.4 (53.7), tail 44.3-48.0 
(45.8), culmen from base 13.0-14.1 (13.5), tarsus 18.2-18.7 (18.5) 
mm. 

Resident. The Scrub Greenlet is found through the Tropical Zone 
lowlands of the Pacific slope of Panama up to 450 m elevation, but 
mainly lower, from the Costa Rican boundary through Chiriqui (D1- 
vala, Bugaba, Concepcion, David, base of Cerro Flores, San Félix), 
Veraguas (Sona, Rio de Jésus, Santa Fé), Herrera (El Rincon), Los 
Santos (Monagre), Coclé (Nata), Panama (Nueva Gorgona, Panama, 
Chico, Puerto San Antonio on lower Rio Bayano) through the Canal 
Zone to the Caribbean side (Corozal, Miraflores, Mount Hope, Juan 
Mina, Empire, Gatun), and adjacent eastern Colon (Colon, Porto- 
belo). Ridgely (in litt.) saw 1 in a garden adjacent to the Panamonte 
Hotel in Boquete, Chiriqui (ca. 1000 m) on January 17, 1974; the bird 
was a full adult with a pale eye and pinkish bill. This race is also found 
in southwestern Costa Rica. 

In fresh plumage these greenlets are bright yellow-green above and 
even brighter yellow below, but these vivid colors fade quickly, espe- 
cially on the upper surface where they become decidedly grayish. The 
breast and abdomen also become paler. The general appearance then is 
surprisingly different. It is this that has led to an attempt to separate 
another race in western Chiriqui under the name pallescens (M. E. 
McClellan Davidson, Proc. Biol. Soc. Washington, vol. 45, 1932, p. 
168). At first it appeared that this was valid, but as more specimens 
were assembled it became evident that the birds are uniform in char- 
acter throughout the mainland of Panama. Those of Los Santos and 
of Colon are similar in every way to a long series of others. 

This greenlet is most frequent as an inhabitant of growths of low 
scrub, but finds second growth and more open forest also to its liking. 
The thorny thickets and other fairly open stands of low trees and 
bushes of the savanna areas are especially suited to its needs and here 
if is fairly common. On the eastern side of the Azuero Peninsula I 


240 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


recorded it only near the sea in northeastern Los Santos but it is prob- 
able that it ranges near the coast down to Punta Mala since the entire 
region contains suitable habitat. Without much doubt it has increased 
in abundance on the Atlantic slope of the Canal Zone and in adjacent 
Colon through the clearing of heavy forest and the appearance of low 
second growth that covers neglected or abandoned fields. In the early 
period it was a rare bird in this section. McLeannan seems to have 
taken few (possibly only 3 in addition to the type) as in his day the 
Caribbean slope where he worked was mainly heavily forested. It thus 
seems reasonable to establish the restricted type locality as Frijoles. 
Galbraith, who assisted in the collecting when the type was secured, 
noted that it was rare. 

These greenlets are found in the higher thickets and in low trees, 
more rarely in higher tree tops. Their movements usually are slow 
and deliberate, with birds pausing quietly to look about. The head often 
appears large in proportion to the size of the body, so that in profile 
they sometimes suggest a miniature peppershrike. Occasionally they 
are more active, somewhat like little warblers, flying up to examine the 
underside of leaves, and then may be more difficult to follow. The pale- 
colored eye is the outstanding character that identifies them when they 
are near at hand. Eisenmann (Condor, 1962, p. 507) describes a distinct 
whistle, “usually of two notes, repeated interminably, which varies 
somewhat individually or seasonally. Usually to my ear the call is 
towheé, towheé, towheé, towheé, repeated unchangingly from five to 
twenty or more times, without pause.” 

It is usual to find this greenlet in pairs, less frequently in groups of 
3 or 4 individuals that I have supposed might be family parties. Breed- 
ing pairs were collected in southern Veraguas. On August 16, 1974, 
Eisenmann saw a fully grown fledgling being fed in a residential sub- 
urb of Panama city. I have no other information regarding their nest- 
ing. Eisenmann (op. cit.) says that in the Pacific Coast of Panama 
“this species is ecologically and geographically sympatric with both 
Hylophilus aurantufrons and Vireo flavoviridis. It seems more tolerant 
of humid conditions than aurantiufrons and less so than flavoviridis, 
as it occurs in scrubby cleared areas of the rainy Caribbean slope, but in 
my experience on the mainland, it avoids forest borders. Generally 
does not forage as high as flavoviridis.” 

The stomach of a bird collected by E. A. Goldman on May 24, 1911, 
at Portobelo, Colén, contained remains of more than 4 spiders 75%, 
a small hymenopteran fragment 3%, jaw and abdomen of a small black 
earwig 4%, a blatted ootheca 3%, bits of elytra of a metallic appearing 


FAMILY VIREONIDAE 241 


buprestid 3%, 25 seeds of Melastomaceae 5%. Another collected Ieb- 
ruary 14, 1912, at Empire contained one hairy and one smooth cater- 
pillar skin, both small 20%, 10 seeds of Panicum sp. and other vegeta- 
ble fragments 80%. Of several taken by Strauch (Bull. Brit. Orn. 
Club, 1977, p. 64), 5 males ranged in weight from 11.3 to 13.0 g, and 6 
females from 11.4 to 13.7 g. 


HYLOPHILUS FLAVIPES XUTHUS Wetmore 


Hylophilus flavipes xuthus Wetmore, 1957, Smiths. Misc. Coll., 1943 (9), p. 85. 
(Isla Coiba, Panama.) 


Characters.—Undersurface much darker, more buffy (instead of 
yellow), sides darker; darker green above than H. f. viridiflavus; bill 
heavier. Immature darker above and below, with wash of ochraceous- 
orange on abdomen. 

Measurements.—Males (7 from Isla Coiba), wing 55.8-58.3 (57.5), 
tail 47.0-50.9 (48.4), culmen from base 14.3-15.0 (14.7), tarsus 17.9- 
10 186) mam. 

Females (6 from Isla Coiba), wing 54.4-56.7 (55.9), tail 45.2-49.3 
(47.8), culmen from base 14.6-16.1 (15.1), tarsus 18.2-19.4 (18.6) 
mm. 

Resident. On Isla Coiba these were among the more common of the 
small birds, though seen infrequently because they ranged among 
screening leaves and creepers. During my visit from January 6 to 
February 6, 1956, they were found in the scrub growths back of the 
beaches and at the borders of mangroves, and came also into the brushy 
rastrojo of old fields, habitats similar to those inhabited by other forms 
of the species in mainland localities. On Coiba, I found that they also 
lived in the high crown of the inland forest, though it was near the end 
of my stay before I verified this, owing to the difficulty to detecting 
small birds in such situations. They move actively among the leaves 
and twigs, almost as quickly as wood warblers, and when seen often 
appear very close at hand. Occasionally I found them feeding on small 
drupes of fruiting trees. The yellowish-white iris of the adult birds is 
often apparent as they climb and hop among the smaller branches. The 
darker colored immature birds have dark eyes. The song, given in low 
tones, usually has three similar notes, swee, swee, swee, which are easily 
imitated by whistling. 

The birds of Coiba are much darker than the nearby mainland race 
and the bill is heavier at the base than in viridiflavus in addition to being 
slightly longer. 


242 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Eisenmann and E. S. Morton in October 1965 and Ridgely in early 
April 1976 found them also within tall forest areas of Coiba, but only 
in lower growth. The notes heard were essentially as on the mainland. 


Family PARULIDAE: Wood Warblers, Reinitas 


Forty-eight species of wood warblers have been found in Panama. 
This total includes 34 North American migrants, mostly from the east- 
ern half of the continent, that winter in Panama or pass through on 
their way north or south. Some are extremely common, especially in 
the periods from September to early November and again in March 
and April, while others are known from only a single or very few rec- 
ords and are still accorded hypothetical status. One species, the Yellow 
Warbler (Dendroica petechia) has both migrant and resident popula- 
tions. Many of the migrant and wintering species join mixed flocks of 
endemic birds, but some others become relatively sedentary while in 
Panama and have been shown to maintain a territory. There is a strong 
correlation between winter territoriality and retention of the “spring” 
plumage; the warblers that have a less boldly marked basic plumage are 
the ones that usually join mixed-species flocks. Many of the North 
American species that are primarily insectivorous in the breeding sea- 
son consume considerable amounts of fruit in Panama. 

Most of the resident wood warblers of Panama are found in the high- 
lands: the three species of Geothlypis resemble G. trichas of North 
America in behavior; a Vermivora and a Parula are highly arboreal, 
usually foraging high in trees; the redstarts of the genus Myioborus 
usually range in the lower levels of the forest; and the 5 species of 
Basileuterus are found in forest undergrowth. Two wood warblers in- 
habit the borders of streams and rivers rather like the waterthrushes 
(Seirus) of North America: Basileuterus fulvicauda is widespread 
in the lowlands and foothills, and the Zeledonia, Zeledonia coronata, is 
rare and local in forest streams in the highlands of Chiriqui and Ver- 
aguas. Although Zeledonia has in the past been placed in its own family, 
Sibley (Postilla, no. 125, 1968, pp. 1-12) and Raikow (Bull. Carnegie 
Mus. Nat. Hist. no. 7, 1978, p. 28) have shown that it is an aberrant 
parulid. 


KEY TO SPECIES OF PARULIDAE 


1. Throat or most of undersurface bright orange or yellow.............. 2 
Undersurface without bright orange or yellow... .: 2). 401.5925 22 see ol 
Z. throat black, \eray,,or dusky... Sack diese Mia es ee 3 


10. 


le 


WZ 


US) 


14. 


Sy 


16. 


7. 


18. 


FAMILY PARULIDAE 


guiconkenoulblack vonay NOt CiISIyin, .& o.5) sca nist Mele Melee nels: Pter sll Wes Maunoihen 
Throat color similar to that of side of face 
Side of face bright yellow. 


Cy 


male Hooded Warbler, Wilsonia citrina. p. 
PMIMUen eA ENeI S's Ot CCMISML. (ori! di sale Wrclinerces Mabe ta mest nie sc islsxeulsseurwtale ore 
Central tail feathers black, outer ones white. 
Slate-throated Redstart, Myioborus miniatus. p. 
PNM MeVe- mines 11 COmpleter OL /a Senta ji enenisraciieetsic enin ec = cae siete 
Complete white eye-ring. 
Connecticut Warbler, Oporornis agillts. p. 
Wing length minus tail length equal 10-18 (13) mm. 
Mourning Warbler, Oporornis philadelphia. p. 
Wing length minus tail length equals 2-10 (6.5) mm. 
MacGillivray’s Warbler, Oporormts tolmiet tolmiei. p. 
Whidersiitkace streaked with brownior blacker sianse cs sce. cl tate cel te 
Wiicrsiinbace | WitOUt iSsineakine 0. Sos). cecghin sume a tie acme aie sha eusrdie een 
Se Rao AChy Ore GIS ae. Asa sie nee Be ee MOI al tao 6 elas 
Streaking bright reddish brown. 

Yellow Warbler, Dendroica petechia. p. 
eres RAE MAVVIENIHE MOU UIT ce4 he, otras eiste teal iced a res eane a Alle Ves ke a ae, 
LENE). SUNN SIRO Be SE Wee RBG RSERRN er ahr CME tee ai ep ny a Ge UG vest te 
HO ilete rateMICOMLEM LGR VUINLCS Med cite eo srk lt ovata tue egal eco aia Sool eeleie sas 
Undertail coverts yellow. 

Palm Warbler, Dendroica palmarum palmarum. p. 
Upper surface blue-gray. 
Yellow-throated Warbler, Dendroica dominica. p. 
Upper surface black or grayish brown. 
Blackburnian Warbler, Dendroica fusca. p. 
LESLEURAT SEIN GY Sa ci eRe a ROE TTI ec eh ean ahh I 
Entire upper surface gray. 
Canada Warbler, Wilsonia canadensis. p. 
Side of face gray or black, large white patch on nearly all tail feathers. 
Magnolia Warbler, Dendroica magnolia. p. 
Side of face bright brown, small amount of white on outer tail feathers. 
male Cape May Warbler, Dendroica tigrina, in alternate plumage. p. 
WipemcuneacesbltevOreslate Chay. s iaucis cs 2 eke nein oe wc couege tities ev alhce Glace 
lipper suniaceroreenSiOrvyellowislt). 2402 40 selene es ome oe ca ws 
NFO SHOnAMnGERSmacenVeMOw eins ce Ware ee nee oe Lil's Gikcac a Ue 
Throat and breast bright orange, rest of undersurface gray. 
Flame-throated Warbler, ’ermivora gutturalts. p. 
Blackish band across breast. 
Collared Redstart, Myioborus torquatus. p. 
Undersurface without band. 
Tropical Parula, Parula pitiayumi. p. 
RNAI Ce COMETS TUM Malta ce Clete Pusey cain Catal Senter (eho feet tye a) ena fe) oe ph Atay RS LG 
Middle and greater wing coverts tipped white. 
Blue-winged Warbler, Vermivora pinus. p. 
mitine ELOmilye LOWISh NDEOW NOs Sthealeqassy miele. scree site. rieren le = 
CrowmwwAthout streaking yellowish. On bROwale yale a see eae Clee 18 Rle es 


244 


19. 


20. 


7. Ih 


Ge. 


23: 


24. 


Za: 


26. 


27. 


28. 


“a. 


30. 


ar 


32. 


JD: 


34. 


BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Crown brown or streaked. ise isi Mee es 
Crown bright yellowish, contrasting with rest of upper surface. 


Prothonotary Warbler, Protonotaria citrea. p. 


Crown brown. 


Rufuous-capped Warbler, Basileuterus rufifrons. p. 


Crown with two wide black stripes. 


Golden-crowned Warbler, Basileuterus culicivorous godmani. p. 


Forecrown yellow, contrasting with rest of crown.................0-+-- 
Forecrown green, gray, or black. 2....6 040.4... eee ee 
Outer tail feathers with large white patches. 


female Hooded Warbler, Wilsonia citrina. p. 


No white on tail feathers. 


Wilson’s Warbler, Wilsonia pusilla. p. 


Large, 150 mm or more. 


Yellow-breasted Chat, Icter1a virens virens. p- 


Smaller, 135 mmvuor less: f.) 2.00: 020002000030. se eeeee 
Superciliarly indistinct\or absent: io.) 5.2 
Prominent yellow superciliary. 


Kentucky Warbler, Oporornis formosus. Pp. 


Black lorésuor maski) iis cdn ee EO eee 
Side of face yellowish; gray, or light brown. ...2.2): 2): 22: s0eeeeeeeee 
Black extensive on side of faces...) 060 
Black confined to lores and area beneath eye. 


Gray-crowned Yellowthroat, Geothlypis poliocephala ridgwayt. p. 


Most of crown gray. 


male Masked Yellowthroat, Geothlypis aequinoctialis chiriquensts. p. 


Grown not (extensively. gray. 6. sis ce Ue ok eke on os 
Forecrown black, bordered by grayish white band. 


male Common Yellowthroat, Geothlypis trichas. p. 


Most of crown black, becoming green on hind crown. 


male Olive-crowned Yellowthroat, Geothlypis semiflava bairdi. p. 


Entire undersurface ‘yellow... 5:0)..024 snd. eee ces oss se 
Belly buff. 


female Common Yellowthroat, Geothlypis trichas. p. 


Much of crown and side of head gray. 
female Masked Yellowthroat, Geothlypis aequinoctials chiriquensis. p. 
Crown and side of head yellowish green. 


female Olive-crowned Yellowthroat, Geothlypis semiflava bairdi. p. 


Basal half of tail. feathers light orange, yellow, or buff... . 42-3 aoe eeeeee 
Tail feathers entirely dark or with white patches extending to distal end. . 
Basal half of all innermost tail feathers orange or yellow. 


American Redstart, Setophaga ruticilla. p. 


Basal half of all tail feathers buff. 


Buff-rumped Warbler, Basileuterus fulvicauda. p. 


Throat entirely black, brown), OF QUAY si. cyanide tes laisse ae 
Throat light coloredor strealed:;.\..) 48) s\sencs dies setsow an toe ee ee 
Throat and upper breast contrasting with rest of undersurface.......... 
Entire undersurface dark gray. 


Zeledonia, Zeledoma coronata. p. 


299 


296 
28 


294 


298 
30 


294 


296 


298 


306 


330 


35. 


36. 


7. 


38. 


31), 


40. 


Al. 


42. 


45. 


46. 


47. 


48. 
49. 


50. 


FAMILY PARULIDAE 245 


Bact DLG ley Or wie mpayicaiacsiacts Pa creal Aree RAMA AEN aia Michi it hoi ye 36 
Throat brown. 
male Bay-breasted Warbler, Dendroica castanea in alternate plumage. p. 280 
cnn ORECTIMOIVELLOWT Sayer, \s lent ian) CRON NEM MN (nous CE 93 37 
Upper surface blue. 
male Black-throated Blue Warbler, Dendroica caerulescens. p. 271 
Wing coverts tipped bright yellow. 
Golden-winged Warbler, Vermivora chrysoptera. p. 250 
Wing coverts tipped white. 
adult male Black-throated Green Warbler, Dendroica virens. p. 275 


Grove bolaily. Strealcedy ics yy ik hee Me RENCE Call Oy al 39 
Crowmeavariciine styeales, on unstreaked jc. Yuu an nD gi 43 
Giown streaks black with brown, butt, .Or eneem.. sees. ul lees le eee ee. 40 


Crown streaks black and white. 
Blalk-and-white Warbler, Mmotilta varia. p. 246 
Mdemstimiace | UNSiEAKeM ees ic Lys ein oe ee) RPG MEME EEC) eG 41 
Undersurface white with dark streaks on breast and flanks. 
Ovenbird, Seiurus aurocapillus. p. 284 
Canmaienow ia stripe DU VOL CeEMISIHA vs skies yall s ulalslahere cists uiei eis ele 42 
Central crown stripe reddish brown. 
Black-cheeked Warbler, Basileuterus melanogenys. p. 318 
Undersurface buff and white. 
Worm-eating Warbler, Helmitheros vermivorus. p. 249 
Undersurface greenish and yellow. 
Three-striped Warbler, Basileuterus tristriatus. p. 313 


PEO MpemNcticnaGce: WNnSteaked |) ii 4).2). baie ue ee eatin eek Wet as 44 
Wippenrsstinmace with imme streaks. 05,0) sl si GaN ACO 51 
Yellow or yellow-green rump contrasting with rest of upper surface...... 45 
INIGi COLMUESIS SV ALSA TABI OY OLN MAL argo SEU a es eT Ct Ee 46 


Rest of upper surface brownish, throat white. 
female or immature Yellow-rumped Warbler, 
Dendroica coronata coronata. p. 271 
Rest of upper surface greenish, throat finely streaked. - 
female or basic plumage male Cape May Warbler, Dendroica tigrina. p. 271 
Whidenstirtace heavily: streaked Nasi ius amuses Meaah ML ba 47 
Wider supraceiwithy little on iO) stealer jie vee en eyacen ie ies Mien Nh Bik 48 
Superciliary pure white, undersurface whitish, culmen of male 
14.4-16.9 (15.5), of female 14.2-16.4 (15.5) mm. 
Louisiana Waterthrush, Seiurus motacilla. p. 288 
Superciliary buffy or dull white, undersurface usually tinged pale yellow, 
culmen of male 12.0-14.1 (13.2), of female 12.0-13.9 (13.1) mm. 
Northern Waterthrush, Seturus noveboracensis. p. 286 
NVincRcoverts promimently tipped white: usa ats c:. ute ia cade ole meet 49 
Nimo COVERS ENOL TIPPECs WiIItee ya Mune ic telson mere ARAL UM LN ous Uk 50 
Upper surface yellowish green. 
female or immature Black-throated Green Warbler, Dendroica virens. p. 275 
Base of primaries white. 
female Black-throated Blue Warbler, Dendroica caerulescens. p. 271 
Base of primaries unmarked. 
Tennessee Warbler, Vermivora peregrina. p. 252 


246 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


51. Crown bright green ‘or with patch of yellow...) 299-0 eee eee 52 
Crown: dull green, blue,sor) blacks. 245) jl) eee 53 
52. Rump yellow. 
adult male Yellow-rumped Warbler, Dendroica coronata coronata. p, 271 
Rump streaked green and black. 
Chestnut-sided Warbler, Dendroica pensylvanica, p. 278 
53. Crown blueor: black). 2.5.5. aes LS ee eee 54 
Crowit) greens pices neh So ee 55 
54. Crown blue. 
adult male Cerulean Warbler, Dendroica cerulea. p. 275 
Crown black. 
male Blackpoll Warbler, Dendroica striata in alternate plumage. p. 282 
55. Undertail coverts white. 
female or male in basic plumage Blackpoll Warbler, 
Dendroica striata. p. 282 


Undertail coverts buffy. 
female or male in basic plumage Bay-breasted Warbler, 
Dendroica castanea. p. 280 


MNIOTILTA VARIA (Linnaeus), Black-and-white Warbler, 
Reinita Trepedora 


Ficure 22 
Motacilla varia Linnaeus, 1766, Syst. Nat., ed. 12, vol. 1, p. 333. (Hispaniola.) 


Small; striped black and white. 

Description.—Length 108-125 mm. Adult male, crown black with 
white stripe down center; back and scapulars black, streaked white; 
rump and upper tail coverts black; middle and greater wing coverts 
black, tipped white, forming two wing bars; remiges dusky black, ter- 
tials edged with white; tail black with white patch on inner web of 
outer three pairs of rectrices; superciliary white; side of face black; 
moustachial stripe white; throat black; sides of breast, flanks, and un- 
dertail coverts striped black, rest of undersurface white. 

Adult female, like male but with throat, side of face and most of un- 
dersurface white. 

Measurements.—Males (10 from eastern North America, taken in 
April), wing 67.0-72.0 (69.4), tail 45.9-49.7 (47.4), culmen from base 
11.1-13.7 (12.5), tarsus 15.2-17.2 (16.4) mm. 

Females (10 from eastern North America, taken in May), wing 
62.0-68.0 (65.1), tail 45.6-48.6 (46.8), culmen from base 11.8-13.3 
(12.6), tarsus 16.3-17.7 (16.9) mm. 

Migrant and winter visitor from the north. The winter range is 
from the southern United States through the West Indies, Middle 
America, and northwestern South America to Venezuela and Ecuador. 


FAMILY PARULIDAE 247 


Fairly common, arrives in Panama as early as August 24 (Arbib and 
Loetscher, Auk, 1935, p. 327), with some remaining until early April 
(April 4, 1911, Lion Hill, Canal Zone, specimen; April 5, 1976, Pipe- 
line Road, Canal Zone, Ridgely); commonest as a migrant in September 
through mid-October and March. The Black-and-white Warbler is 
found throughout the Republic in winter, although less numerous east 
of the Canal Zone; it is commonest in the lowlands, but I have seen it 
as high as 1560 m at Cerro Punta, Chiriqui. It has also been found on 
Taboga and Urava Islands and on San José on the Pearl Islands; the 
British Museum of Natural History has a specimen collected October 
25, 1924, at sea in the Gulf of Panama. 


Ficure 22.—Black-and-white Warbler, Reinita Trepedora, Mniotilta varia, male. 


As a migrant and winter visitor the Black-and-white Warbler be- 
haves just as it does in the north, climbing the trunks and main branches 
of trees in search of insects. Buskirk et al. (Auk, 1972, p. 619) found 
that at Cerro Punta Black-and-white Warblers will join the local inter- 
specific flocks of insectivorous birds, but will not travel with them 
for any distance. I have also seen them with other migrant warblers. 
Skutch (A Naturalist in Costa Rica, 1971, p. 104) found that in Costa 


248 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Rica Black-and-white Warblers maintain territories during the win- 
ter and may sing while driving away intruders. In Panama an in- 
dividual banded by H. Loftin September 7, 1963, at Almirante, Bocas 
del Toro was recaptured there on October 7, 1963, and January 8, 1964. 
One collected by Strauch (Bull. Brit. Orn. Club, 1977, p. 64) weighed 
HOsSi 


PROTONOTARIA CITREA (Boddaert): Prothonotary Warbler, 
Reinita Protonotaria 


Motacilla citrea Boddaert, Tabl. Planch. enlum., 1783, p. 44. (Louisiana.) 


Small; bright orange-yellow with gray wings and tail. 

Description.—Length 111-125 mm. Adult male, head, nape, and en- 
tire undersurface except undertail coverts bright orange-yellow; back 
yellowish olive-green, changing to bluish gray on rump and upper tail 
coverts; scapulars yellowish olive-green; wing coverts bluish gray; 
alula black with outer web broadly edged white; remiges black with 
outer web edged bluish gray and inner web edged white; rectrices gray 
with white patch on inner web of all but central pair; undertail coverts 
white; edge of wing yellow; underwing coverts white. 

Adult female, like male, but crown and hindneck as well as upper 
back yellowish olive-green; yellow on undersurface less orange. 

Measurements.—Males (10 from Panama), wing 67.1-73.8 (70.8), 
tail 39.6-50.8 (45.9), culmen from base 13.1-14.7 (13.8), tarsus 15.6- 
S26) (793) eaoaaaae 

Females (10 from Panama), wing 64.0-70.1 (67.0), tail 42.3-51.1 
(45.0), culmen from base 12.9-15.3 (13.8), tarsus 14.6-18.7 (17.5) 
mm. 

Migrant and winter visitor from the north. It winters from southern 
Mexico to northern Colombia and Venezuela, and casually to Ecuador, 
the Guianas, and West Indies. Found throughout the Tropical Zone in 
wet localities, common in the lowlands in mangrove swamps and in 
thickets along streams, mainly those of larger size, but when in mi- 
gration these warblers may appear along the smaller tributaries in the 
mountain foothills to at least 300 m (Quebrada Cauchero, Serrania de 
Majé, Province of Panama, March 6, 1950). Most arrive from the 
north in September—the earliest report is August 1 (Eisenmann, 
Smiths. Misc. Coll., vol. 117, no. 5, 1952, p. 50)—and are present until 
the middle of March, a few remaining later. Ridgely has found it as 
late as March 20, in 1979 at El Volcan, Chiriqui. The Prothonotary 
Warbler has been found on Isla Coiba, Taboga, Taboguilla, and Urava, 
and on San José, Rey, and Pacheca Islands in the Perlas group. 


FAMILY PARULIDAE 249 


The main migration periods are September to mid-October and early 
February to mid-March; during the spring migration period nearly all 
the birds I collected were heavy with fat. For the most part, this species 
leaves Panama relatively early. In areas where they are common in 
mid-winter, such as Juan Diaz, Province of Panama, they have already 
decreased markedly by the beginning of March: in 1976 Ridgely found 
more than 40 there on January 24, but only 2 on March 10. They may 
remain more numerous later westward; Ridgely found more than 30 in 
mangroves at Estero Rico, western Chiriqui, on March 6, 1976. 

During migration, Prothonotary Warblers range from mangrove 
thickets to at least 30 m up in trees in swampy woods. On March 3, 
1948, I saw dozens feeding in flowering trees at E] Rincon, Herrera. 
They feed commonly in Erythrina trees around Tocumen and else- 
where in the Canal Zone during January and February; often they are 
with Tennessee Warblers, orioles, and other species. On February 5, 
1955, at Isla Taboga, at dusk over 100 descended on a group of trees; 
they were noisy and easy to approach, but were gone the next morning 
(F. O. Chapelle in litt. to Eisenmann). The weights of 3 collected by 
Burton (Bull. Brit. Orn. Club, 1975, p. 85) range from 12.4 to 13.5 g. 


HELMITHEROS VERMIVORUS (Gmelin): Worm-eating Warbler, 
Reinita Gusanera 


Motacilla vermivora Gmelin, Syst. Nat., vol. 1, pt. 2, 1789, p. 951. (Philadelphia, 
Pennsylvania. ) 


Small; crown striped buff and black; rest of upper surface olive- 
green; undersurface cream buff. 

Description.—Length 112-131 mm. Adult (sexes alike), crown 
striped light olive-buff and black; rest of upper surface, tail, and wing 
coverts olive-green; black line through eye; superciliary pale buff; side 
of head and upper breast olive-buff; flanks grayish olive; throat and 
rest of undersurface buffy white; underwing coverts buffy white. 

Measurements.—Males (10 from eastern North America, taken in 
April), wing 67.0-73.0 (69.6), tail 43.7-55.4 (49.4), culmen from 
base 13.9-15.6 (15.2), tarsus 17.0-19.3 (18.0) mm. 

Females (10 from eastern North America, taken in April and May), 
wing 64.1-70.5 (66.9), tail 43.2-53.7 (48.0), culmen from base 13.2- 
15.8 (14.4), tarsus 16.8-18.6 (17.6) mm. 

Winter visitor from the north. Uncommon but regular in west, rarer 
eastward; recorded east to the Canal Zone (Barro Colorado Island) 
and eastern Province of Panama (Puerto San Antonio, on the Rio 
Bayano), the southern limit of its winter range. Several of the records 


250 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


are from the Subtropical Zone forests of Chiriqui, at Santa Clara, Bar- 
riles, and Pena Blanca, above Boquete. Arcé obtained it in Veraguas at 
Santa Fé and Chitra. I have 1 from El Uracillo, on the Caribbean slope 
of Coclé, and there is 1 in the British Museum collected by McLeannan 
along the Panama Railroad. Specimens have been taken from October 
12 (1938, Pena Blanca, Chiriqui) to March 16 (1954, Santa Clara, 
Chiriqui; 1927, Puerto San Antonio, Panama) and April 8 (1963, 
Almirante, by Olson). Netting at Almirante extends early and late 
dates from September 24, 1964, to April 17, 1965 (D. Hicks), and an 
extreme date of April 23,1963 (lorttin, Carib. Journ senya asa 
154). 

A Worm-eating Warbler banded October 21, 1963, at Almirante, 
Bocas del Toro, was recaptured there on January 8, 1964 (Loftin et al., 
Bird-Banding, 1966, p. 41). Another individual caught in a mist net 
on the Chiva Chiva Road in the Canal Zone on December 15, 1968, 
weighed 12.9 g. 


VERMIVORA CHRYSOPTERA (Linnaeus): Golden-winged Warbler, 
Reinita Alidorada 


Motacilla chrysoptera Linnaeus, Syst. Nat., ed. 12, vol. 1, 1766, p. 333. (Philadel- 
phia, Pennsylvania. ) 


Small; crown yellow; yellow patch on wings; rest of upper surface 
gray; face and throat black or gray; undersurface white. 

Description.—Length 107-120 mm. Adult male, superciliary and 
cheek patch white; crown bright yellow; a large black patch covering 
orbital area and ear coverts; stripe on side of face white; most of mid- 
dle and greater wing coverts, forming a large yellow patch; tail gray 
with extensive white patch on inner web of outer three pairs of rec- 
trices; throat black; sides of breast and flanks light gray, rest of under- 
surface white; underwing coverts white. 

Adult female, like male, but crown sometimes tinged with olive- 
green; rest of upper surface varying from entirely gray to heavily suf- 
fused with olive-green; areas of face and throat that are black in male 
are gray in female. 

Measurements.—Males (10 from Panama), wing 55.5-65.8 (61.3), 
tail 44.4-51.7 (48.4), culmen from base 12.2-13.8 (12.9), tarsus 15.6- 
17.9 (16.7) mm: 

Females (10 from Panama, Costa Rica, and Colombia), wing 57.0- 
61.2 (58.4), tail 42.7-51.9 (46.6), culmen from base 11.8-13.9 (12.4), 
tarsus 15.5-18.6 (17.1) mm. 

Migrant and winter visitor from the north. It winters from Guate- 


FAMILY PARULIDAE 251 


mala to central Colombia and northern Venezuela. Fairly common in 
the Tropical and Subtropical Zones, primarily recorded from the moun- 
tains of Chiriqui, where it has been collected as high as 2310 m ( Bangs, 
Proc. New England Zool. Club, vol. 3, 1902, p. 63), and in the Canal 
Zone (Barro Colorado Island, Lion Hill, Juan Mina). In Bocas del 
Toro the Golden-winged Warbler has been collected at Cocoplum 
(Chapman, Auk, 1931, p. 121) and at Almirante (Loftin and Olson, 
Carib. Journ. Sci., vol. 3, no. 4, 1963, pp. 194-195). On March 10, 
1961, I took a male with no fat at Tacarcuna Village, Darién. Ridgely 
(in htt.) found as many as 12 above Santa Fé, Veraguas, on January 4, 
1974. 

In Panama this species is most often noted while on migration, from 
mid-September to early November and late March to mid-April. The 
extreme dates are September 19 (Eisenmann, Smiths. Misc. Coll., vol. 
iMiArnes, 1952) p) 5b) and) April 18 (Lottinand Olson} op./cit.).. 

Buskirk et al. (Auk, 1972, p. 619), studying mixed species flocks at 
Cerro Punta, Chiriqui, found that Golden-winged Warblers were long- 
term followers in these flocks, although rarely were more than 1 or 2 
individuals part of a flock. Ridgely notes that it usually occurs in 
forest canopy or at its edge and almost always with mixed flocks. This 
species often feeds by probing in clusters of dead leaves hanging from 
branches. 

There seem to be no reports from the drier parts of the Pacific slope 
lowlands from western Province of Panama, southern Coclé, and the 
Azuero Peninsula, and few east of the Canal Zone. Eisenmann and 
others have observed it occasionally on Cerro Campana and Cerro 
Azul, and he and Morton saw 1 on Isla Coiba on October 9, 1964. 


VERMIVORA PINUS (Linnaeus): Blue-winged Warbler, Reinita Aliazul 
Certhia pinus Linnaeus, 1766, Syst. Nat., ed. 12, 1, p. 187. (Philadelphia, Penn- 


sylvania. ) 

Small; crown and undersurface bright yellow; most of upper sur- 
face bright olive-green; wings and tail gray. 

Description.—Length 103-115 mm. Adult male, crown lemon yel- 
low; black line through eye; rest of upper surface bright olive-green; 
wings gray with middle and greater coverts tipped white, forming two 
wing bars; tail gray with large white patches on inner web of outer 
three pairs of rectrices; undertail coverts white; rest of undersurface 
lemon yellow; underwing coverts white. 

Adult female, like male, but crown duller, often greenish; line 
through eye paler; undersurface sometimes tinged with green. 


252 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


Measurements.—Males (10 from New Jersey, taken in May), wing 
58.3-62.5 (60.1), tail 43.3-47.9 (46.5), culmen from base 11.3-12.5 
(11.9), tarsus 16.0-18.0 (17.1) mm. 

Females (10 from New Jersey, taken in May), wing 55.0-59.0 
(56.6), tail 42.2-46.6 (44.3), culmen from base 10.6-12.7 (11.7), tar- 
sus 15.9-17.8 (16.8) mm. 

Winter visitor from the north. This species normally winters from 
southern Mexico south to Nicaragua; in Panama it is a rare visitant 
infrequently recorded in the western Caribbean lowlands, both slopes 
of the Canal Zone, where it was first recorded on December 30, 1942, 
at Cano Saddle on the northwestern shore of Gatun Lake (Imhof, Auk, 
1950, p. 256), and in the Province of Panama as far east as Chepo, 
where it has been collected once, March 7, 1927 (Griscom, Auk, 1933, 
p. 306). The Smithsonian has a specimen collected at Almirante, Bocas 
del Toro, on January 17, 1961. The Blue-winged Warbler has been 
found in Panama between mid-October and late March. An individual 
of the hybrid form known as “Brewster’s Warbler” was seen at Vol- 
can Lakes, Chiriqui, on January 13, 1977, by A. Greensmith (Toucan, 
vol. 4, no. 2, 1977, p.6). N. G. Smith (im litt. to Eisenmann) noted a 
bird with “no mask, whitish below with yellow breast patch” on the Siri 
Grande, Bocas del Toro, on January 8, 1964. 


VERMIVORA PEREGRINA (Wilson): Tennessee Warbler, 
Reinita Peregrina 


Syliva peregrina Wilson, Amer. Orn., vol. 3, 1811, p. 83, pl. 25, fig. 2. (Banks of 
the Cumberland River, Tennessee. ) 


Small; upper surface mostly bright olive-green; undersurface mostly 
white. 

Description.—Length 103-113 mm. Adult (sexes alike), crown and 
hindneck bright olive-green, sometimes partially replaced by gray in 
birds from February or later; rest of upper surface bright olive-green, 
more yellowish on rump and upper tail coverts; tail blackish, edged 
bright olive-green; superciliary white or yellowish white; undersurface 
white or suffused to greater or lesser degree with greenish yellow; un- 
dertail coverts white; underwing coverts white. 

Measurements.—Males (10 from Panama), wing 60.5-65.2 (62.8), 
tail 40.3-45.3 (42.8), culmen from base 10.3-11.9 (11.2), tarsus 15.4- 
18.1 (16.7) mm. 

Females (10 from Panama), wing 57.2-64.1 (62.0), tail 37.8-46.9 
(41.2), culmen from base 10.6-12.3 (11.4), tarsus 14.1-17.2 (16.3) 


mm. 


FAMILY PARULIDAE Ze 


Migrant and winter visitor from the north. Winters from southern 
Mexico to Colombia and Venezuela. Very common in the Tropical and 
Subtropical Zones on both Atlantic and Pacific slopes of Panama east 
through Chiriqui (especially in the mountains), Veraguas, and Bocas 
del Toro; fairly common farther east. The species has been recorded 
on Isla Coiba, Taboga and Taboguilla islands, and San José, Saboga, 
and FE] Rey (San Miguel) islands in the Perlas Archipelago. Aldrich 
pndnBole (Scient. Publ. Cleveland Mus. Nat. Hist., vol. 7, 1937, p. 
123) found the Tennessee Warbler to be the most abundant of all the 
North American migrants on the western slope of the Azuero Penin- 
sula. This species is more tolerant of conspecifics than are most winter- 
ing warblers. Early on the morning of March 19, 1957, on the Rio Oria, 
in southern Los Santos, I flushed a migrant flock of more than 100 
from the edge of a stream where they had come down to drink. They 
flew in a fairly close formation, like goldfinches, and swarmed through 
the trees. 

In addition to insects, nectar is an important food of Tennessee War- 
blers in winter. Tramer and Kemp (Auk, 1979, pp. 186-187) describe 
how in Costa Rica individual birds defend from conspecifics as large a 
section as they can of a flowering Erythrina tree. Morton (pers. 
comm.) has observed similar behavior in Panama. Skutch (pers. 
comm. to Ridgely) has found the Tennessee Warbler the only parulid 
to come to his bird feeder. When foraging for insects, the birds are 
found from open lawns to forest canopy, usually in groups. 

Tennessee Warblers are present in Panama from mid-September 
until late April. Ridgely and J. Pujals saw 1 at Ft. Sherman, Canal 
Zone, on May 1, 1976. Six were banded at the Gorgas Memorial Lab- 
oratory Station at Almirante, Bocas del Toro, between April 18-24, 
1963 (Loftin and Olson, Carib. Journ. Sci., vol. 3, no. 4, 1963, p. 195). 
Recaptures of birds at Almirante show that some spend several weeks 
or possibly the entire winter there; an individual banded on October 21, 
1963, was recaptured there on February 25, 1964 (Loftin et al., Bird- 
Banding, 1966, p. 41). Another banded at Almirante on October 21, 
1964, was recaptured there the following year on October 20, 1965 
(Loftin et al., Bird-Banding, 1967, p. 151). One collected by Burton 
Chul Brit. Orn. Club, 1975, p..85), at Jaqué, Darien; weighed) 7.5 ¢g: 

The record by Murphy (Fish and Wildlife Service Spec. Sci. Rep 
Fisheries, no. 279 [mimeo], 1958, p. 110) of the Nashville Warbler, 
Vermivora ruficapilla, near Coiba and off the Gulf of Panama in No- 
vember 1956 refers to the Tennessee Warbler, the notes having been 
placed inadvertantly under the wrong heading. 


254 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


[VERMIVORA RUFICAPILLA (Wilson): Nashville Warbler, 
Reinita Cabecigris 


Sylvia ruficapilla Wilson, 1811, Amer. Ornith., 3, p. 120, pl. 27, fig. 3. (Near 

Nashville, Tennessee.) 

An individual of this species was reported seen at Santa Clara, Chiri- 
qui, on January 2, 1980, by P. E. Donohue and J. Van Os (Ridgely, in 
hitt.). The normal winter range is from southern Texas and Mexico 
to Guatemala.] 


VERMIVORA GUTTURALIS (Cabanis): Flame-throated Warbler, 
Reinita Gargantirroja 


Compsothlypis gutturalis Cabanis, 1860, Journ. f. Ornith., 8, p. 329. (Irazu, Costa 

Rica.) 

Small; throat and breast bright orange; upper surface gray with 
black triangular patch on back. 

Description.—Length 106-116 mm. Adult (sexes alike), entire up- 
per surface slate gray except for black triangular patch on upper back; 
primaries blackish with outer web narrowly edged slate gray; secon- 
daries blackish on inner web, slate gray on outer web; rectrices blackish 
with outer web edged slate gray; lores and side of face blackish; throat 
and breast rich orange to cadmium orange (sometimes almost flame 
scarlet) flanks light gray; rest of undersurface white. 

Juvenile, upper surface olive-gray with blackish triangular patch on 
back; wings and tail dusky, with narrow buffy tips to upper wing co- 
verts; lores and face dusky; narrow, buff postocular stripe; chin and 
throat orange-yellow, fading to buff on abdomen. 

A female taken March 2, 1965, at Volcan de Chiriqui, Chiriqui, had 
the iris dark brown; base of gonys and lower margin of mandibular 
rami dull orange-yellow; rest of bill black; tarsus and toes dull mouse 
brown; claws somewhat darker brown. 

Measurements.—Males (10 from Chiriqui and Costa Rica), wing 
61.4-65.1 (63.2), tail 43.6-52.0 (47.6), culmen from base 12.3-13.8 
(13.0), tarsus 16.2-18.1 (17.3) mm. 

Females (10 from Chiriqui and Costa Rica), wing 57.0-65.0 (60.4), 
tail 42.8-50.7 (46.2), culmen from base 10.6-13.1 (12.4), tarsus 16.0- 
17.6 (16.8) mm. 

Resident. Fairly common in forest and forest borders of western 
Chiriqui (Volcan de Chiriqui, Cerro Punta, and Boquete), where it 
has been collected between 1560 and 3090 m (Blake, Fieldiana: Zool., 
vol. 36, no. 5, 1958, pp. 557-558). Of the specimens collected by Mon- 
niche at the elevations given by Blake, “well over half’? were taken be- 


FAMILY PARULIDAE 255 


low 1800 m. This species is also found in the mountains of Costa Rica. 
Four specimens from January and February 1902 labeled by Batty, 
Brava Island, Chiriqui, and Cébaco Island, Veraguas, are certainly er- 
roneously marked (Eisenmann, Auk, 1950, p. 364). 

The Flame-throated Warbler feeds in typical warbler fashion, moy- 
ing quickly through the branches of shrubs and trees in search of in- 
sects; it ranges from treetops to lower levels at woodland edges and 
shrubby clearings. It often hangs down from pendant leaves that it 
examines carefully. Buskirk et al. (Auk, 1972, p. 619), studying inter- 
specific foraging flocks at Cerro Punta, considered this species a long- 
term follower in such flocks, although rarely were more than 2 indi- 
viduals part of any flock. Ridgely (1976, p. 292) describes the song as 
“a weak dry buzz, zeeeeoo.” Mr. and Mrs. Rodman Ward (in litt. to 
Eisenmann) reported seeing this species on April 3, 1959, at 2300 m 
on Cerro Punta and watched 1 as it uttered a buzz zeeeeee-u (slightly 
inflected at the end), and heard (without seeing the singer) a very high- 
pitched buzzy tsip-i-tsip-i-tsip-i-tee. Eisenmann has heard only call 
notes such as tsip; tick, tick; or dick dick. 

Nothing is recorded about the nesting habits of this bird in Panama, 
but Skutch (Nutt. Orn. Club, no. 7, 1967, pp. 137-143) has studied it 
in detail in Costa Rica, where he found them nesting from mid-March 
through June. Half the nests Skutch found were well up in trees, be- 
tween 6 and 25 m from the ground, and half were in niches on grassy 
banks. All the nests, whether terrestrial or arboreal, though not roofed, 
were well covered by growing vegetation, either by dense grass or moss 
or by epiphytes or broad-leafed bromeliads on trees. The nest, built 
entirely by the female, is made primarily of green moss and liverworts; 
some fine plant fibers and horsehair may be used in the lining, but even 
here green moss predominates. One nest measured by Skutch was 8 
cm in height by 10 or 12 cm in diameter; the cavity was 5 cm in diameter 
by 3 cm deep. In every case that Skutch could determine, the clutch 
was 2 eggs, these vary in pattern from unspotted white to faintly and 
finely sprinkled with brown over the entire surface. Measurements of 
4 eggs were from 16.2 12.9 to 19.0 13.1 mm. 

Incubation is done exclusively by the female and takes approximately 
16 days. The young hatch with orange-pink skin and long but sparse 
gray down. Their mouth lining is yellow on the marginal regions and 
pink or pale red in the more central and deeper regions, unlike the 
mouth linings of most passerine nestlings, which are uniform yellow or 
yellow-orange. The young are fed small green caterpillars by both 
parents and leave the nest at approximately 13 days. 


256 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


[PARULA AMERICANA (Linnaeus ): Northern Parula, Reinita Nortena 


Parus americanus Linnaeus, 1758, Syst. Nat., ed. 10, 1, p. 190. (South Carolina.) 


There are two recent sightings of this species, both from Galeta 
Island, Canal Zone: one was found there by J. Pujals and others on 
January 21, 1976, the other was seen on March 4, 1977, by Pujals and 
S. Stokes (Ridgely, im litt.). Both appeared to be adult males. Puyjals 
reports that on March 4, 1977, when he saw the Northern Parula, he 
also saw a Palm Warbler (Dendroica palmarum), a species whose 
tropical winter range is also mainly in the West Indies. The Northern 
Parula normally winters in Florida, the West Indies, Mexico, and 
Central America south to Nicaragua.| 


PARULA PITIAYUMI (Vieillot): Tropical Parula, Reinita Mariquita 
Sylvia pitiayumi Vieillot, 1817, Nouv. Dict. Hist. Nat., ed., 11, p. 276. (Paraguay. ) 


Very small; upper surface dull, dark blue; most of undersurface 
rich yellow with throat and breast brownish orange. 

Description.—Length 88-110 mm. Adult (sexes alike), lores, sides 
of head dull indigo blue to bluish slate gray, except for triangular olive- 
green patch in center of back; extent, if any, of white tipping on mid- 
dle and greater wing coverts varying geographically; rectrices black 
with outer webs edged blue and broad subterminal patch of white on 
the inner web of two outermost pairs; throat and breast deep orange- 
ochraceous; sides and flanks blue; rest of undersurface to undertail 
coverts rich lemon yellow; undertail coverts white; underwing coverts 
white. 

Parula pitiayumi is one of the most widely distributed of the tropical 
warblers, although its range is discontinuous and the bird is often ab- 
sent from humid lowland forest areas; it breeds from southern Texas 
south through Mexico, Central America, Panama, and in South Amer- 
ica from Colombia to the Guianas, Peru, Bolivia, Uruguay, and north- 
ern Argentina. Fourteen races are recognized, of which 3 occur in 
Panama—-speciosa in lower highlands and foothills of western Panama, 
cirrha on Isla Coiba off Veraguas, and nana in the lowlands and lower 
foothills of eastern Province of Panama and Darién. In Panama, the 
Tropical Parula is a bird of tall forest, but elsewhere, in southern Texas 
and eastern Venezuela and the chaco of southern South America, for 
example, it inhabits low, dry woodlands, and in the Cauca Valley of 
Colombia it lives in semi-arid cultivated valleys with scattered trees. It 
is a very active feeder, reminiscent of the Northern Parula. It often 
feeds in flowering trees. 1 have sometimes seen them with flocks of 


FAMILY PARULIDAE 257 


migrant warblers. The song, in Chiriqui “a buzzy trill, typically tsip- 
tsip-tsip-tsip-tsip-tsrrrrrrrrrrip, with variations” (Ridgely, 1976, p. 
292), is delivered throughout the day, and through many months of the 
nonbreeding season, when most other birds are silent. 


PARULA PITIAYUMI SPECIOSA (Ridgway) 
Compsothlypis pitiayumi speciosa Ridgway, 1902, Auk, 19, p. 69. (Boquete, Chiri- 
qui, Panama.) 

Characters.—Sides of face blackish; lower throat and breast orange- 
ochraceous; abdomen yellow; one short white wing bar or spot formed 
by tipping on the greater wing coverts. 

A male collected March 22, 1965, at El Volcan, Chiriqui, had the iris 
dark brown, cutting edge of maxilla and mandible light yellow; rest of 
maxilla black; tarsus dull brown; toes dull honey yellow; claws dull 
grayish brown. 

Measurements.—Males (10 from Chiriqui and Costa Rica), wing 
49.0-55.0 (52.0), tail 36.7-41.0 (38.5), culmen from base 9.6-11.5 
(10.5), tarsus 15.0-16.9 (15.9) mm. 

Females (9 from Chiriqui), wing 48.0-49.9 (49.0), tail 34.9-38.8 
(36.9), culmen from base 9.6-11.1 (10.3, average of 8), tarsus 14.6- 
16.4 (15.4) mm. 

Resident. Fairly common in forests in the lower highlands of 
Chiriqui and in the foothills of Veraguas and Herrera. W. W. Brown, 
Jr. (Bangs, Proc. New England Zool. Club, vol. 3, 1902, p. 63) col- 
lected it at Boquete, Chiriqui between 1000 and 1350 m, and Monniche 
(Blake, Fieldiana: Zool., vol. 36, no. 5, 1958, p. 558) found it at vari- 
ous localities on the Volcan de Chiriqui between 1560 and 1920 m. An 
unusual record is the specimen taken by J. W. Batty on Brava Island, 
off the coast of Chiriqui on January 28, 1902, but like many Batty speci- 
mens this is probably erroneously labeled (Eisenmann, Auk, 1950, p. 
364). In Veraguas and Herrera it has been recorded from 450 to 600 
m; Arcé collected it in Veraguas at Colobre (Salvin and Godman, Biol. 
Centr.-Amer., Aves, vol. 1, (pt. 6), 1880, p. 120). 

Nothing has been published on this species’ nesting behavior in Pan- 
ama, but Skutch (Nutt. Orn. Club, no. 7, 1967, pp. 143-146) has found 
a nest of P. p. speciosa in Costa Rica. On May 6, 1964, he observed a 
female carrying material to a cushion of green moss about 10 m up ona 
nearly vertical branch of a tree in a grove near heavy forest; the nest 
was within the cushion of moss, with a round entrance on the side. 
Skutch never saw the eggs from this nest, but mentions that in Trini- 
dad they have “an almost glossless white ground .. . [and are] spotted 


258 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 
and blotched with deep or pale chestnut, forming a more or less distinct 
cap on the broader end.”’ Three weeks after discovery, the nest con- 
tained 3 well-feathered nestlings in the entrance; both parents were 
bringing them green caterpillars, a small, green, grasshopper-like in- 
sect, and tiny white protein corpuscles from a Cecro pia tree. 

Ridgely (im litt.) reports that on June 7, 1969, he found a nest near 
FE] Volcan. It was on the underside of a moss-covered limb, “burrowed 
into the moss,” about 10 m up in a roadside tree, in a patch of humid 
woodland, 5.1 km below E1 Volcan, on the highway. Some grass and 
twigs projected from the nest. 


PARULA PITIAYUMI CIRRHA Wetmore 


Parula pitiayumi cirrha Wetmore, 1957, Smiths. Misc. Coll., 134 (9), p. 88. (isla 
Coiba, Panama.) 


Characters.—More deeply colored above and below than P. p. spe- 
ciosa; green patch on back very small; sides of face blackish-blue; 
throat lemon chrome; lower breast and abdomen orange, continuous 
with chest color; conspicuous white spot on greater wing coverts. 

Mcasurements.—Males (4 from Isla Coiba), wing 52.4-58.5 (55.8), 
tail 39.2-41.7 (40.8), culmen from base 12.1-12.8 (12.5), tarsus 15.7- 
17.8 (16.8) mm. 

Females (2 from Isla Coiba), wing 53.5-54.7 (54.1), tail 39.2-39.8 
(39.5), culmen from base 11.7-12.6 (12.2), tarsus 16.7-16.8 (16.8) 
mm. 

Resident on Isla Coiba. On three occasions during my visit to Isla 
Coiba in 1966 I saw pairs of these birds moving quickly among leafly 
twigs in the high branches of tall forest trees, so high above the ground 
that they were barely within gunshot. Except for their restless move- 
ments they would never have been detected in the shaded light of these 
forest haunts. Only occasionally did they appear briefly in silhouette 
against some tiny opening that led to the open sky. Probably they were 
fairly common, as the forest cover of the entire great island was suited 
to their needs. 

During a short stay (October 8-10, 1965) Eisenmann and E. S. 
Morton found the Coiba birds common in all woodland, usually in pairs 
between 1 and 8 m up in trees, on the border of woods or partly cut- 
over forest. None were singing. On Ridgely’s visit (April 9-14, 1976) 
he found the birds singing very persistently and saw them at all levels 
in the forest. 


FAMILY PARULIDAE 259 


PARULA PITIAYUMI NANA (Griscom) 


Compsothlypis pitiayumi nana Griscom, 1927, Amer. Mus. Novit., no. 282, p. 8. 
(Cape Garachiné, eastern Panama [ = Darién].) 


Characters.—Smaller than P. p. speciosa; sides of face same color 
as crown, lighter than in P. p. speciosa.; two white wing bars formed 
by tipping on middle and greater wing coverts. 

Measurements.—Male (1 from Darién), wing 49.0, tail 36.8, cul- 
men from base 10.6, tarsus 15.0 mm. 

Female (1 from Darien, the type), wing 47.2, tail 31.7, culmen 
from base 10.4, tarsus 14.2 mm. 

Resident. Local, but in some areas common in the lowlands and 
lower foothills of eastern Province of Panama (“hills just east of 
Chepo, on road from Platanares to Jésus Maria, eastward; common 
in Bayano River valley” (Ridgely, 1976, p. 292) and in Darién, where 
it is known from Garachiné, the type locality, and from Cana, where 
R. R. Benson collected a “breeding”’ female in the spring or summer of 
1923 (Griscom, Bull. Mus. Comp. Zool., vol. 69, 1929, p. 183). Gris- 
com (Bull. Mus. Comp. Zool., vol. 78, 1935, p. 367) assigned this race 
to the “Arid Tropical Zone,” although Cana is in fact humid. Nothing 
is recorded of its breeding in Panama or in northwest Colombia, where 
it also occurs. In mid-January 1975 Ridgely twice observed pairs feed- 
ing fledged young on middle slopes of Cerro Quia (500-670 m). 
Ridgely also found this species at Cana, where they were uncommon, 
but has not seen it in the lowlands of Darién. 

The birds from the Bayano River basin have been assumed to be 
nana, although specimens have not been collected. 


DENDROICA PETECHIA (Linnaeus): Yellow Warbler, 
Reinita Mangletera 


FIGURE 23 


Motacilla petechia Linnaeus, 1766, Syst. Nat., ed. 12, 1, p. 334. (Barbados.) 


Small; mostly bright yellow, more greenish on upper surface; male 
of resident races with entire head and upper breast reddish brown, 
chest, and sides streaked reddish brown; males of migrant races have 
reddish brown only as streaks on breast, chest, and sides. 

Description.—Length 110-117 mm. Adult male of resident races, 
entire head, throat, and breast reddish brown (in jubaris mixed with 
yellow); rest of upper surface yellowish green, slightly lighter on 
rump; wing feathers dusky blackish, edged lemon yellow; on remiges 


260 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


edge of inner web also lemon yellow; rectrices dusky blackish, edged 
yellow on outer web with much of inner web also yellow; rest of under- 
surface and underwing coverts yellow, with fine reddish brown streaks 
on chest and flanks. Adult male of migrant races, similar, but brown 
of head and throat replaced with yellow, bright on crown; reddish 
brown streaking begins at breast. 


fj 


wiki Sd ] 


Mee VYWUVEZ zw 
~ 7p lie 


Ficure 23.—Yellow Warbler, Reinita Mangletera, Dendroica petechia, a male of 
the erithachorides (Mangrove Warbler) group. 


FAMILY PARULIDAE 261 


Adult female, like male, but paler, duller yellow throughout, and 
reddish brown areas absent or fainter and reduced. 

Immature, like female, but paler. 

The Yellow Warbler is the only member of its family represented in 

Panama by both migrant and resident races. Dendroica petechia com- 
prises three groups of subspecies that were once considered separate 
species: the yellow-headed aestiva group (Yellow Warbler), which 
is mainland North American in distribution and chiefly migratory, the 
chestnut-capped petechia group (Golden Warbler) of the West Indies, 
most of coastal Venezuela and Cozumel Island, and the chestnut- 
hooded erithachorides group (Mangrove Warbler) of both coasts of 
Central and South America to Peru on the Pacific side and to the Para- 
guana Peninsula of Venezuela on the Atlantic side. Except when they 
occur on relatively small islands, birds of the petechia and erithacho- 
rides groups are usually confined to coastal mangrove swamps. Mi- 
grants are widely distributed in the lowlands in semi-open areas and 
gardens, as well as in mangroves. 

Recent works (e.g., Lowery and Monroe, in Peters, Check-list Birds 
World, vol. 14, 1968, pp. 14-20) include the erithachorides and aestiva 
groups in the species petechia. Large series of specimens from Pan- 
ama and Colombia recently studied by Olson (Proc. Biol. Soc. Wash., 
vol. 93, no. 2, 1980, pp. 473-480) corroborate this treatment. The 
Yellow Warblers on the Pacific Coast from Panama to Peru exhibit a 
gradation in plumage characters from birds of the chestnut-hooded 
erithachorides group to those resembling the chestnut-capped petechia 
group. The subspecies on the Galapagos and Cocos Island is presum- 
ably derived from the erithachorides group, but is convergently similar 
to the West Indian birds. 

The song, behavior, and diet of the resident races of the Yellow War- 
bler are similar to those of North American forms, except that, as 
noted, the former are usually confined to mangroves. The stomach of 
a resident individual collected in the Canal Zone by E. A. Goldman on 
May 30, 1911, contained 2 blunt caterpillar jaws 5%, 3 entire spider 
eggs parasitized by a hymenopteran, and fragments of others 35%, a 
tipulid 12%, head of a hymenopteran 1%, moth remains 1%, 14 heads 
and other remains of ants 30%, a small chrysomelid 1%, weevil re- 
mains 1%, 4 beetles ( Nitidulidae, Colastus?) 5%, other coleoptera 9%. 
Apparently these birds breed somewhat irregularly: of 2 adult males I 
collected at La Honda, Los Santos, in March 1948, 1 was singing and 
in full breeding condition, while the other, like it in plumage, had the 
testes small. 


262 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


The migrant races of the Yellow Warbler are very common in the 
lowlands of both slopes and on the islands off the Pacific Coast, as well 
as inland. They are found in a variety of habitats from forest border 
to mangrove to gardens and grassy areas at the edge of taller vegetation. 
They often deliver chip or tsip notes, but I have never heard a migrant 
or wintering Yellow Warbler sing in Panama; Skutch (A Naturalist 
in Costa Rica, 1971, p. 104), however, has found individuals maintain- 
ing a winter territory from which they may sing while driving away in- 
truders. A male of one of the migrant races collected by E. A. Gold- 
man at Cana, Darién on March 19, 1912, contained 1 Leptostylus sp. 
25%, 2 small cerambycids 2%, 1 elatrid 2%, 3 minute calandrids 4%, 
another beetle 2%, bits of a membracid 2%, caterpillar remains 25%, 
an arachnid and 1 egg 3%, 12 ants 30%, 2 other Hymenoptera 5%. 

In the Canal Zone, Eisenmann considers the migratory Yellow War- 
bler, during the months of the northern winter, the fourth most com- 
mon parulid, exceeded in numbers only by the Bay-breasted, Chestnut- 
sided, and Tennessee Warblers. 

H. von Wedel (Peters, Bull. Mus. Comp. Zool., vol. 71, 1931, p. 333) 
collected a migrant female at Almirante, Bocas del Toro, on August 13, 
1928, and L. L. Jewel (Auk, 1913, p. 428) recorded it at Gatun, Canal 
Zone, on May 12, 1912. Most migrants do not arrive before late Au- 
gust, and depart by late April. A Yellow Warbler banded at Almirante 
on November 11, 1962, was recaptured there 9 weeks later, on January 
9, 1963 (Loftin, Bird-Banding, 1963, p. 220). 

Mr. M. Ralph Browning has very kindly identified the migrant Yel- 
low Warblers from Panama now in the Smithsonian collections. 


DENDROICA PETECHIA AMNICOLA Batchelder 


Dendroica aestiva amnicola Batchelder, 1918, Proc. New England Zool. Club, 6, 
p. 82. (Curslet, Newfoundland. ) 


Characters.—“Similar to D. a. aestiva, but stated to differ in the male 
sex by darker green, less yellowish—between warbler green and sul- 
phine yellow—back; more restricted and duller yellow of the forehead; 
narrower as well as duller, citron yellow rather than strontian yellow, 
edges to the remiges; and on average less richly colored underparts 
with darker chestnut streaks; female duskier, less yellowish above. .. . 
distinguished [from rubiginosa] by the yellow forehead contrasting 
with the green of the back.” (Hellmayr, Field Mus. Nat. Hist. Zool. 
Ser.) vol, 13, pt,/81935).p) 563)) 

The race amnicola breeds from north-central Alaska to central Labra- 


FAMILY PARULIDAE 263 


dor and south to central Alaska, northeastern British Columbia and 
across Canada to south-central Quebec, Newfoundland, and the Magda- 
len Islands; it winters from southern Baja California and central 
Mexico south to Peru, Colombia, and French Guiana. 

The 17 specimens of this race in the Smithsonian Institution collected 
in Panama were taken on the Pacific slope from San Félix, Chiriqui, to 
La Jagua in eastern Province of Panama, and on the Caribbean slope 
from Almirante, Bocas del Toro, to Juan Mina, Rio Chagres, Canal 
Zone. I also have specimens from Isla Cébaco and Isla Coiba, off the 
Pacific Coast of Veraguas. M. EF. Davidson collected a female at Puerto 
Armuelles, Chiriqui, on November 27, 1929 (California Academy of 
Science no. 32798). 


DENDROICA PETECHIA RUBIGINOSA (Pallas) 


Motacilla rubiginosa Pallas, 1811, Zoogr. Rosso-Asiat., 1, p. 496. (Kodiak Island, 
Alaska.) 


Characters.—Similar to aestiva, but slightly smaller and much duller 
in color. Adult male darker and dull olive-green above, the pileum con- 
color with the back or else becoming slightly more yellowish on fore- 
head (very rarely distinctly yellowish on forehead and forepart of 
crown); wing edgings less conspicuous, mostly yellowish olive-green, 
sometimes inclining to yellow on greater coverts. Adult female darker 
and duller olive greenish above, duller yellow below. (Ridgway, U.S. 
Nat. Mus., Bull. 50, pt. 2, 1902, p. 514) 

This race breeds on the west coast of North America from southern 
Alaska to western British Columbia. It winters from Mexico to Pan- 
ama. Five specimens from Panama were taken at Pedasi, Los Santos, 
and La Jagua and Chiman, Province of Panama, on the Pacific slope, 
and at Juan Mina, Rio Chagres, Canal Zone, and Puerto Obaldia, San 
Blas, on the Caribbean slope. All were females or immature males. 
M. E. Davidson took a female at Puerto Armuelles, Chiriqui, on No- 
vember 27, 1929 (California Academy of Science no. 32799) and Gris- 
com identified specimens from Almirante, Bocas del Toro, as belonging 
EO this race ( Peters, Bull. Mus. Comp, Zool., vol. 71, 1931, p. 333). 


DENDROICA PETECHIA AESTIVA (Gmelin) 
Motacilla aestiva Gmelin, 1789, Syst. Nat., 1 (2), p. 996. (City of Quebec, Canada.) 
Characters —Upper surface more decidedly olive-green, the upper 


tail coverts with less yellow; streaks on chest and sides much broader 
than in other races. 


264 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


The race aestiva breeds from southeastern Alberta across Canada to 
Prince Edward Island and Nova Scotia and south to central southern 
Montana, central Oklahoma, northern Arkansas, central Alabama, 
central Georgia, and central South Carolina. It winters from central 
Mexico to Peru and northern Brazil. 

Nineteen specimens of aestiva in the Smithsonian collection from 
Panama were all from the Pacific slope, from Puerto Armuelles, Chiri- 
qui, through Los Santos and Herrera to El Llano on the Rio Bayano, 
Province of Panama, and El Real, Darién. Four were taken on Isla 
Coiba, Veraguas, and 1 on Isla de Canfas, in the Pearl Islands Archi- 
pelago. One at Buena Vista, Chiriqui, was at 690 m. A male in the 
British Museum (no. 85.3.8.387) was taken by Arcé in 1868 at Calove- 
vora, Veraguas, on the Caribbean slope. Of the several taken in March, 
only 1 male showed extensive streaking. 

Ralph Browning has found that 2 Yellow Warblers from Panama 
in the Smithsonian collection are from a distinctive population from 
Nebraska that is characterized by more heavy streaking than in aestiva, 
morcoml, or sonorana, more intense yellow on the undersurface, and, 
on the upper surface, it is usually paler than all other races except 
sonorana. This population may merit description as a new subspecies. 
The 2 Panama specimens are USNM no. 386476, a male taken at 
Jaqué, Darién, on April 8, 1946, and USNM no. 457705, a female 
from Ft. Clayton, Canal Zone, taken September 22, 1953. 


DENDROICA PETECHIA MORCOMI Coale 
Dendroica aestiva morcom Coale, 1887, Bull. Ridgway Ornith. Club, no. 2, p. 82. 
(Fort Bridger, Utah [ = Wyoming].) 
Dendroica aestiva brewsteri Grinnell, 1903, Condor, 5, p. 72. (Palo Alto, Cali- 
fornia.) 
Dendroica aestiva ineditus Phillips, 1911, Auk, 28, p. 85. (Matamoros, Tamauli- 
pas, Mexico.) 


Characters.—Slightly larger than aestiva, duller (less yellow) green 
coloration; breast streaking heavy. 

This race breeds from western Washington, central British Colum- 
bia, and eastern Montana south to southern California and northwest- 
ern Baja California across to northwestern Texas. It winters from 
Baja California to Ecuador, northern Colombia, Venezuela, and French 
Guiana. Three specimens from Panama are all males: USNM no. 
461264 from San Félix, Chiriqui, February 21, 1956; no. 533282 from 
Punta Balsa, Chiriqui, February 25, 1966; and no. 462376, from Pedasi, 
Los Santos, March 9, 1957. 


FAMILY PARULIDAE 265 


Griscom (Bull. Mus. Comp. Zool., vol. 78, 1935, p. 367) recorded the 
subspecies sonorana from Veraguas and the Canal Zone but there are 
no specimens from Panama identified as sonorana in either the Mu- 
seum of Comparative Zoology (fide R. A. Paynter, Jr.) or the Ameri- 
can Museum of Natural History (fide W. E. Lanyon). Nor are there 
any specimens of this subspecies in the extensive Smithsonian holdings 
of migrant D. petechia from Panama. D. a. sonorana breeds in the 
southwestern United States and northern Mexico and reputedly win- 
ters as far south as Colombia and Ecuador, so it might be expected to 

-occur in Panama. This remains to be confirmed, however. 


DENDROICA PETECHIA AEQUATORIALIS Sundevall 


Dendroica petechia aequatorialis Sundevall, 1870, Ofv. K. Sv. Vet.-Akad. Forh., 
26 (1869), p. 609. (Panama City.) 


Characters.—Male with crown, cheeks, and throat uniform brown, 
but of a lighter shade than in aithocorys, upper breast streaked with 
yellow. 

A male collected at Chico, Province of Panama, on March 24, 1961, 
had the iris wood brown; maxilla fuscous; base of mandibular rami 
across base of gonys dull buffy brown, rest of mandible dull neutral 
gray; tarsus and toes dull honey yellow; claws mouse brown. 

Measurements.—Males (10 from Province of Panama), wing 59.8- 
63.5 (62.5), tail 43.6-48.8 (47.1), culmen from base 11.1-13.0 (12.0), 
tarsus 17.3-18.8 (18.2) mm. 

Females (10 from Province of Panama), wing 58.5-62.5 (59.6), 
tail 39.3-49.3 (45.6), culmen from base 11.5-13.3 (12.2), tarsus 17.1- 
18.7 (17.9) mm. 

Resident. Common but somewhat local on the Pacific Coast of the 
Province of Panama and on the Pearl Islands. On the mainland it is 
usually found in mangroves, but occasionally in other vegetation ad- 
jacent to salt water. They are, for example, widespread on the forti- 
fied islands of Ft. Amador, in scrub and low dry woods. In 1944, how- 
ever, on San José and Pedro Gonzalez islands in the Pearl Archipelago, 
I found that Yellow Warblers were distributed widely through the 
higher forest growths. This is probably due to the absence of compet- 
ing small insectivores such as are found on the mainland. On San José 
I recorded them away from the shore, mainly in the taller trees near or 
over water, but on Pedro Gonzalez they were common over the forested 
hills far from any such habitat. Because of the heavy vegetation they 
were more evident along the shores, as elsewhere they remained con- 


266 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


cealed in the leaves 20 to 25 m from the ground. In various places 
small trees with thin, light green leaves grew along the shore, often 
projecting out from rocky ledges or sandy banks over the tide line. 
These were favorite haunts, as were open-branched trees growing at 
the tops of higher, steeper bluffs. A male I collected on San José on 
February 19 was in breeding condition. 

As Thayer and Bangs (Bull. Mus. Comp. Zool., vol. 46, 1905, pp. 
155-156) noted, males from the Pearl Islands sometimes have the yel- 
low portions of the plumage replaced with orange; there are 2 such 
males from San José in the Smithsonian collection. Some of the adult 
males from Majé, on the mainland, show traces of yellow in the cheeks, 
perhaps an indication of intergradation with the race jubaris. 

The songs of aequatorialis in Panama City (heard at a mangrove islet 
near Old Panama on July 16, 1950) reminded Eisenmann of that of 
aestiva in New York, and were of two types, both rather sweet: chee, 
chee, chee, wichoo-eécha, with variants, and a more ringing swee, swee, 
swee, chéw, chichoo cheéoo. Calls were chip (or chit) and a fast chit- 
chitchitchitchit. 


DENDROICA PETECHIA AITHOCORYS Olson 


Dendroica petechia atthocorys Olson, 1980, Proc. Biol. Soc. Wash., 93(2), pp. 
474-475. (5 miles east of La Honda, near Los Santos, Los Santos Province, 
Panama. ) 

Characters.—Compared to aequatorialis, male with brown portions 
darker, upper breast not streaked with yellow; females of both races 
variable, but aithocorys generally with more streaking below than 
aequatorialis. 

A male collected at Puerto Aguadulce, Coclé, on March 11, 1962, had 
the iris light dull reddish brown; maxilla and tip of mandible fuscous- 
black; base of mandible dull neutral gray; tarsus and toes light buffy 
brown; claws neutral gray. A female taken the same day had the cutting 
edge of maxilla and mandible dull light neutral gray; rest of maxilla 
fuscous-black; tarsus and toes and claws dull brown. Another male 
collected at Aguadulce on January 18, 1963, had the iris dark brown; 
maxilla fuscous-black; basal half of gonys dull flesh color; rest dull 
gray; tarsus and toes dull verona brown; claws dark mouse brown; 
underside of toes dull honey yellow. 

Measurements.—Males (10 from western Panama), wing 59.3-66.0 
(63.6), tail 45.6-51.1 (48.6), culmen from base 12.2-13.9 (12.9), tar- 
sus 17.7-19.3 (18.5) mm. 


FAMILY PARULIDAE 267 


Females (10 from western Panama), wing 58.5-65.5 (61.1), tail 
43.3-48.4 (46.7), culmen from base 11.8-13.6 (12.7), tarsus 16.2-19.6 
(18.1) mm. 

Resident. Common in mangroves on the Pacific Coast of western 
Panama from at least as far west as Puerto Armuelles, Chiriqui, east 
as far as Puerto Aguadulce, Coclé, and on Isla Coiba off the coast of 
Veraguas. On Coiba this species is confined to the coastal mangroves; 
in this it differs from populations on other islands in Panama that are 
often found in tall forest far from salt water. In the Aguadulce area of 
Coclé there is some intergradation with aequatorialis. 

At Puerto Aguadulce on July 4, 1952, Eisenmann saw at least 5 in 
the coastal mangroves. A male, accompanied by a female, sang a very 
fast tit-tit-tit-tit-teé. 

The population on Isla Bolanos and possibly Isla Parida, Chiriqui, 
about 45 km southeast of David, may merit recognition. A male and 2 
females collected on Isla Bolanos are like aithocorys in plumage, but 
are evidently larger (Olson, Proc. Biol. Soc. Wash., vol. 93, no. 1, 1980, 
p. 475). Yellow Warblers are also found on nearby Isla Parida, but 
none have been collected there. 


DENDROICA PETECHIA IGUANAE Olson 


Dendroica petechia iguanae Olson, 1980, Proc. Biol. Soc. Wash., 93(2), pp. 475- 
476. (Isla Iguana, Los Santos Province, Panama.) 


Characters.—Male nearest aithocorys, but brown of head much 
darker, less rufescent; green of dorsum darker, less yellowish, breast 
and particularly the lower belly more densely streaked with chestnut; 
yellow of underparts and tail darker, appearing dingy and greenish. 
Female like aithocorys, but tending to be more heavily streaked, yellow 
of underparts dingier, as in the male. 

Measurements.—Males (7 from Isla Iguana), wing 61.6-67.4 
(65.0), tail 47.2-52.9 (50.1), culmen from base 12.4-14.4 (13.4), tar- 
sus 17.1-19.8 (19.1) mm. 

Females (4 from Isla Iguana), wing 61.5-62.6 (62.0), tail 47.4-49.2 
(47.8), culmen from base 12.2-12.9 (12.5), tarsus 17.8-20.0 (19.0) mm. 

Resident. Found only on Isla Iguana, Los Santos, a small island 
probably less than 1 km? in area, lying approximately 6 km from the 
mainland, about 7 km east-northeast of Pedasi and 18 km north of 
Punta Mala, on the eastern coast of the Azuero Peninsula (7°38/N, 
80°W ). Iguana is the only offshore island along the eastern and south- 


268 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


ern coasts of the Azuero Peninsula. Considering that, with the possible 
exception of birds from Isla Bolafos, mentioned earlier, none of the 
island populations of Yellow Warblers in Panama have differentiated 
from those of the mainland, it is rather surprising to find that those of 
Isla Iguana are so distinct. Even in the field their more greenish colora- 
tion is noticeable. 

The fauna of Isla Iguana is dominated by a great nesting colony of 
Magnificent Frigatebrids (Fregata magnificens) interspersed with 
Brown Pelicans (Pelecanus occidentalis). During my visit in February 
1957, the Yellow Warbler was the only abundant small bird on the is- 
land, where it ranged everywhere, from the tall bunch grass above the 
rocky shore to the scrub forest inland. I saw them hopping about on 
bare rocks above the high-tide mark in early morning, and 15 or 20 
ranged about the outdoor kitchen near the house, where they came out 
familiarly on low perches, as well as higher among the leaves of the 
trees. In the forest I found them often examining tufts of leaves almost 
among the frigatebirds perched in the branches. Many were in pairs, 
and males sang occasionally, though they were not breeding. 


DENDROICA PETECHIA ERITHACHORIDES Baird 


Dendroica erthtachorides [sic] Baird, 1858, in Baird, Cassin, and Lawrence, Rept. 
Expl. Surv. R.R. Pacific, 9, p. 283. (Cartagena, Colombia.) 


Characters.—Males with rufous rather than hazel on head and throat, 
where it is less extensive than in other races; streaks on chest wider. 

Measurements.—Males (8 from Bocas del Toro), wing 63.0-69.5 
(66.5), tail 48.7-56.1 (51.5), culmen from base 11.5-14.7 (13.2), tar- 
sus 19.2-20.7 (19.3) mm. 

Females (6 from Bocas del Toro and Province of Panama), wing 
60.0-65.7 (63.6), tail 44.3-52.9 (49.1), culmen from base 12.0-13.7 
(12.6), tarsus 18.6-21.4 (19.8) mm. 

Resident. Common on the entire Caribbean coast of Panama from 
Almirante, Bocas del Toro, west beyond the border to Magdalena, Co- 
lombia. On Isla Escudo de Veraguas, | found Yellow Warblers scat- 
tered through the taller trees, where they were fairly common, though 
each of the 4 I took on a visit in March 1958 appeared to be alone. On 
Escudo this species was not restricted to the limited growths of man- 
groves found near the sea, as is the case on the mainland, but ranged 
throughout the forest growth, as appears to be its regular habit when 
found on small islands. 

This series from Escudo and Almirante Bay agree fully with type 


FAMILY PARULIDAE 269 


material of this race, which is interesting since specimens from Limon, 
Costa Rica, about 100 km to the north belong to the race D. p. bryanti. 


DENDROICA PETECHIA JUBARIS Olson 
Dendroica petechia jubaris Olson, 1980, Proc. Biol. Soc. Wash., 93(2), p. 478. 

(Nuqui, Dept. Chocd, Colombia, 5°40’N.) 

Characters.—Compared with aequatorialis, male with brown portions 
of plumage lighter, more tawny; throat and chin either suffused with, 
or distinctly streaked with yellow; cheeks and lores suffused with yel- 
lowish, contrasting with the darker crown, so that birds appear dis- 
tinctly capped, as opposed to aequatorialis, in which the entire head is 
brown; belly less streaked with brown, appearing more brightly yellow. 

Measurements.—Males (10 from Darién and Colombia), wing 60.6- 
67.5 (65.0), tail 45.6-50.8 (48.5), culmen from base 11.4-14.0 (12.9), 
tarsus 17.9-19.8 (19.2) mm. 

Females (10 from Darién and Colombia), wing 56.9-64.7 (61.0), 
tail 42.9-50.0 (46.1), culmen from base 11.7-13.6 (12.3), tarsus 17.2- 
ZOMG S!3))) nam, 

Resident. Found in mangroves on the coast of Darien and south 
along the Pacific Coast of Colombia, at least to Buenaventura, Valle del 
Cauca. In March and April of 1946 I found them common at Jaque, 
Darién. Males were singing and in breeding condition. In ascending 
the Rio Jaqué I noted that the Yellow Warblers were found only where 
there were mangroves. 

Eisenmann noted 2 chestnut-headed birds on March 29, 1967, at Isla 
Boca Grande in the upper Gulf of San Miguel, Darién, but most of the 
Yellow Warblers seen there appeared to be of the migratory aestiva 


group. 


DENDROICA MAGNOLIA (Wilson): Magnolia Warbler, 
Reinita Pechirrayada 


Sylvia magnoha Wilson, Amer. Orn., vol. 3, 1811, p. 63, pl. 23, fig. 2. (Fort Adams, 

Mississippi. ) 

Small; crown gray; back olive-green; tail black with large white 
patches; undersurface yellow, sometimes with black streaks. 

Description.—Length 104-117 mm. Adult male, crown and nape 
gray, back light yellowish olive, increasingly marked with black in 
March and April; rump pale lemon yellow; upper tail coverts black; 
primary coverts black; middle and greater secondary coverts tipped 
white, forming a broad white patch; remiges blackish with outer web 
edged light gray, inner web edged white; rectrices black, all but central 


270 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


pair with large white patch on basal half of inner web; side of face 
gray, with lores, forehead, and malar region becoming black by March, 
and lower and upper rims of eye and stripe behind eye beginning to 
appear in February, prominent by March; undertail coverts white, rest 
of undersurface bright yellow with black streaks beginning at breast, 
becoming increasingly prominent after February; underwing coverts 
white. 

Adult female, as male but duller, with black on back fainter or ab- 
sent, black on face absent, and streaks on undersurface fewer or absent. 

Measurements.—Males (10 from the breeding range, taken in June 
and July), wing 57.5-61.8 (59.8), tail 45.2-51.1 (48.4), culmen from 
base 8.4-11.3 (10.0), tarsus 16.1-17.8 (17.1) mm. 

Females (10 from the breeding range, taken in May, June, and July), 
wing 55.1-58.7 (56.8), tail 43.1-48.5 (46.1), culmen from base 8.7-11.3 
(10.3), tarsus 16.1-18.1 (17.5) mm. 

Winter visitor from the north. Uncommon, but regular in very 
small numbers at least on the Caribbean slope. The Magnolia Warbler 
has been recorded on migration in Bocas del Toro at Almirante, where 
2 were banded on October 21, 1962 (Loftin, Carib. Journ. Sci., vol. 3, 
no. 1, 1963, p. 67) and 1 was taken at Cocoplum on November 5, 1927 
(Chapman, Auk, 1931, p. 121). One of the birds banded at the Gorgas 
Memorial Laboratory station at Almirante on October 21, 1972, was re- 
captured there 5 weeks later on December 7 (Loftin, Bird-Banding, 
1963, p. 220). McLeannan forwarded 2 specimens from what became 
the Canal Zone to Salvin (Salvin and Godman, Biol. Centr.-Amer., 
Aves, vol. 1 (pt. 9), 1881, pp. 129-130); these are now in the British 
Museum. Lawrence (Ann. Lyc. Nat. Hist. New York, vol. 7, 1861, p. 
322) records it as taken in the same area by McLeannan and Galbraith, 
and I secured 2 females at Juan Mina in the Canal Zone, on January 12 
and 17,1955. There are several recent sightings from eastern Province 
of Panama as far east as the Bayano River valley (Ridgely, 1976, p. 
293), and I saw 1 at Jaqué, Darién, on March 29, 1946. This species is 
usually found at low levels in second-growth woodland, where it often 
forages with mixed flocks. Ridgely (1m litt.) has found them in man- 
groves. 

In 1977 Magnolia Warblers were recorded on the exceptionally late 
dates of May 27 (Achiote Road, Canal Zone, N. G. Smith) and May 
28 (Majé Island, Bayano River, C. Lowe) (Toucan, vol. 4, no. 7, 1977, 
p. 2). Smith observed that other northern migrants, including Seiwrus 
noveboracensis, Icterus galbula, 3 species of swallow, Coccyzus ameri- 
canus, and C. erythrophthalmus, remained very much later than usual. 


FAMILY PARULIDAE 271 


He attributed this to an exceptionally extended dry season, with “no 
rains until late May.” 


[DENDROICA TIGRINA (Gmelin): Cape May Warbler, Reinita Altigrada 
Motacilla tigrina Gmelin, 1789, Syst. Nat., 1 (2), p. 985. (Canada.) 


Winter visitant from the north. Very rare, known only from recent 
sight records in Chiriqui, Bocas del Toro, and the Canal Zone. Ridgely 
(1976, p. 294) lists 5 sight records: “one at Almirante, Bocas del Toro, 
- on February 2 and 10, 1958 (Wetmore); one at Changuinola, Bocas 
del Toro, on December 7, 1962 (Eisenmann); one below Volcan, west- 
ern Chiriqui, on January 20, 1970 (P. Alden); one bright male at Sum- 
mit Gardens, Canal Zone, on February 2, 1973 (Ridgely, Eisenmann, 
C. Leahy, J. Gwynne); and one male at Volcan, Chiriqui, on January 
15, 1974 (C. Leahy and Ridgely). More recent sightings include 
March 6, 1976, a female at Estero Rico, in western Chiriqui below 
Divala (Ridgely); December 5, 1976, Bayano River Research Station, 
eastern Province of Panama (A. Greensmith, Toucan, vol. 4, no. 2, 
1977, p. 6). Mason (Auk, 1976, p. 168), reviewing the status of this 
species in Middle America, where it is generally considered a vagrant, 
concludes that “the Cape May Warbler is an infrequent winter visitor 
or winter resident in much of Middle America and that its status may 
have changed in the past two decades.’’| 


[DENDROICA CAERULESCENS (Gmelin): Black-throated 
Blue Warbler, Reinita Azulosa 


Motacilla caerulescens Gmelin, 1789, Syst. Nat., 1(2), p. 960. (Santo Domingo.) 


Vagrant. This species is included on the basis of 1 sight record of a 
male near Gattin Locks (besides the French Canal), Canal Zone, on 
January 14, 1973, by D. Engelman, James Smith, N. Gale, and A. 
Ramirez (Ridgely, 1976, p. 294). The normal winter range of D. 
caerulescens is the Bahamas, Greater Antilles, islands of the western 
Caribbean, and, less commonly, northern Central America, and north- 
ern Colombia and northern Venezuela. | 


DENDROICA CORONATA CORONATA (Linnaeus): Yellow-rumped 
Warbler, Reinita Coroniamarilla 


Motacilla coronata Linnaeus, 1766, Syst. Nat., ed. 12, 1, p. 333. (Philadelphia, 
Pennsylvania. ) 


Small; upper surface gray or light brown, with rump and crown 
yellow; undersurface white with yellow on flanks. 


272 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


Description Length 113-131 mm. Adult male, crown and back 
dark gray, with central crown feathers yellow near base, more or less 
exposed, and black streaks on back; rump pale lemon yellow; upper 
tail coverts black, edged gray; lesser wing coverts black tipped gray; 
middle and greater coverts black tipped white, forming two bars; 
remiges dusky blackish with outer webs thinly edged gray; rectrices 
blackish with white spots on inner webs of outer three pairs; lores and 
ring around eye white, sometimes obscured; side of face gray; under- 
surface white, with fine black streaks on breast and flanks, and pale 
lemon yellow patch on upper flank; underwing linings white. 

Adult female, like male, but gray of head and back replaced by gray- 
ish brown. 

Measurements.—Males (10 from Panama and Mexico), wing 69.8- 
751 (72.2), tail 52.1-60.5 (56.0), culmen from base 94712735 tia 
tarsus 17.0-18.6 (18.0) mm. 

Females (10 from Panama), wing 66.0-71.8 (68.6), tail 48.1-56.5 
(53.4), culmen from base 9.2-11.7 (10.4), tarsus 17.3-18.3 (17.9) mm. 

Winter visitor from the north. Fairly common, at times locally 
abundant in the western half of the Republic in the Tropical and lower 
Subtropical Zones; much less common eastward. In Panama the Yel- 
low-rumped Warbler, while found in more open woodlands and in 
shrubbery about houses, also feeds regularly on the ground in pastures, 
cultivated fields, and along the borders of lagoons. In the Chiriqui high- 
lands and about Almirante and Changuinola in Bocas del Toro, I have 
found them spread over lawns, attracted by swarms of leaf-hoppers, 
and elsewhere have noted them in flooded areas or about other bodies of 
water where gnats were abundant. Charles Handley relates that he has 
seen these birds hawking insects 1n the surf at the shore of Isla Basti- 
mentos, Bocas del Toro; the warblers were catching something at the 
edge of the water; then darting up just out of reach of the surf, not 
alighting at any time except on branches of nearby bushes. 

This species has been recorded from November 7, in 1927, at Coco- 
plum, Bocas del Toro (Chapman, Auk, 1931, p. 121) to March 20, 
when in 1958 I saw 1 at the La Jagua Gun Club, east of Pacora in the 
Province of Panama. I have collected many at Santa Clara and El 
Volcan, Chiriqui, Almirante and Changuinola, Bocas del Toro, and 
Cerro Campana, Panama. Loftin and Olson (Carib. Journ. Sci., vol. 3, 
no. 4, 1963, p. 195) mention 1 seen at Tonosi, Los Santos, on March 8, 
1963, and W.W. Brown, Jr., collected a female on San Miguel in the 
Pearl Islands on February 23, 1904 (Thayer and Bangs, Bull. Mus. 


FAMILY PARULIDAE 273 


Comp. Zool., vol. 46, 1905, p. 156). A further eastern record on the 
Caribbean slope is one I saw at Mandinga, in the Comarca de San Blas, 
on January 27, 1957. Ridgely (im litt.) saw a Yellow-rumped War- 
bler in flowering Erythrinas and on the ground in clearings at the Cana, 
Darién, gold mine on February 28 and March 3, 1981. 

From January 13 to February 15, 1958, this was one of the most 
abundant birds in the Almirante-Changuinola region. Following the 
latter date, there was a definite decrease, though the species remained 
common until the end of February. A week later there were few about, 
and the last one was noted on March 10. This abundance seems to have 
been related to severe cold that year in the north, since friends at Al- 
mirante reported to me that none of these warblers were about in the 
winter months of 1958-59. 

As Ficken and Ficken (Bird-Banding, 1966, pp. 273-279) noted, the 
Yellow-rumped Warbler is one of the few members of its family to 
form flocks of conspecifics in the nonbreeding season. At 840 m on 
Cerro Campana I have seen groups of a dozen or more move across the 
slopes, many of them coming up the ravines from the lowlands and ad- 
vancing to the higher levels. 

The only sighting of the race D. c. auduboni in Panama was made by 
R. Forster and Ridgely (1m litt.), who found 1 bird near Volcan, Chiri- 
qui, on February 5-6, 1976. The bird was with a typical nominate 
coronata, so direct comparisons were possible; among other things, 
the prominent pale yellow throat was obvious, as was its distinctive call, 
a fast whit rather than the sharper tchek of nominate coronata. 


[DENDROICA TOWNSENDI (Townsend ): Townsend’s Warbler, 
Reinita de Townsend 


Sylvia townsendi J. K. Townsend (ex. Nuttall MS), 1837, Journ. Acad. Nat. Sci. 
Philadelphia, 7, p. 191. (Forests of Columbia River.) 


Vagrant from the north. This species is included on the basis of 3 
sight reports. A male seen repeatedly at Nueva Suiza in the highlands 
of western Chiriqui on November 19-30, 1967, by T. V. Heatley and 
V.M. Kleen (Ridgely, 1976, p. 295). On March 21, 1979, J. Baird and 
Ridgely (1m litt.) photographed a bright female near Volcan, Chiriqui. 
The bird was in a small grove of exotic cypresses, and was seen there 
again the next day. On March 21 Baird and Ridgely also saw an adult 
male Townsend’s Warbler on the Boquete Trail above Cerro Punta, at 
2100 m. The normal winter range is from California south to Nica- 
ragua. There are also a number of sight reports from Costa Rica.] 


274 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


DENDROICA VIRENS (Gmelin): Black-throated Green Warbler, 
Reinita Verdosa 


Motacilla virens Gmelin, 1789, Syst. Nat., 1(2), p. 985. (Philadelphia, Pennsyl- 


vania. ) 


Small; upper surface green; sides of head yellow; throat black; rest 
of undersurface white. 

Description—Length 104-117 mm. Adult male, crown, back, and 
rump yellowish olive-green, slightly more yellowish on rump; upper 
tail coverts black, broadly edged gray; wings black, with middle and 
greater coverts tipped white, forming two bars, and remiges edged 
gray on outer web and white on inner web; outer two pairs of rectrices 
white with narrow blackish stripe on inner side of outer web, next pair 
blackish with white spot on outer web, remaining pairs blackish or black 
with outer web edged gray; lores, superciliary, and sides of head and 
neck bright yellow; throat, breast, and flanks black, with feathers often 
tipped white until February; rest of undersurface white, with abdomen 
tinged yellow; underwing coverts white. 

Female and immatures, like adult male but green of upper surface 
slightly duller and black on undersurface less extensive. 

Measurements.—Males (10 from the breeding range, taken in May, 
June, and July), wing 60.0-65.8 (62.4), tail 45.0-51.3 (48.2), culmen 
from base 9.3-11.6 (10.8), tarsus 12.8-18.8 (16.5) mm. 

Females (10 from the breeding range, taken in May, June, and July), 
wing 57.0-61.7 (59.6), tail 44.6-51.4 (47.5), culmen from base 9.8-11.3 
(10.6), tarsus 14.5-17.7 (16.6) mm. 

Migrant and winter visitor from the north. Common in the foothills 
and highlands of Chiriqui and Veraguas, less regular but at times still 
common in the hill country of central Panama; in the lowlands it oc- 
curs only asa transient. Monniche (Blake, Fieldiana: Zool, vol. 36, no. 
5, 1958, p. 558) collected this species on the Volcan de Chiriqui between 
1500 and 2700 m; I have found it in this area at the same elevations, 
and at Santa Clara at 1260 m. Farther east, I have found Black- 
throated Green Warblers at El Cope in Coclé, at La Campana (60 m) 
in the Province of Panama, and at Cerro Mali, Darién, where on March 
1, 1964, I shot one at 1260 m on the south fork of the Rio Pucro. In 
the Canal Zone this species is “rather rare” (Ridgely, 1976, p. 295): 
Lawrence (Ann. Lyc. Nat. Hist. New York, vol. 7, 1861, p. 293) re- 
ported that McLeannan collected it on the Atlantic side near the Pan- 
ama Railroad, but Willis and Eisenmann (Smiths. Contrib. Zool. no. 
29, 1979) do not record it from Barro Colorado Island. 

Black-throated Green Warblers have been reported in Panama from 


FAMILY PARULIDAE 275 


mid-September until mid-April; most are found between late Septem- 
ber and late March. Buskirk et al. (Auk, 1972, p. 619), in studying 
mixed-species flocks at Cerro Punta, Chiriqui, considered D. virens a 
“joiner” but not a follower of the flocks led by permanent residents. I 
have often seen them in the flocks of migrant warblers. They are 
usually found in second-growth woodland, borders, and clearings. 


[DENDROICA OCCIDENTALIS (Townsend): Hermit Warbler, 
Reinita Coronada 


: Sylvia occidentalis |.K. Townsend, 1837, Journ. Acad. Nat. Sci. Philadelphia, 7, 
p. 190. (Forests of Columbia River.) 


This species is included on the basis of several recent sight reports. 
An adult male was seen at Nueva Suiza in the highlands of western 
Chiriqui on December 22, 1973, by Dana Gardner and again in the exact 
same locality on January 15, 1974, by Ridgely and C. Leahy (Ridgely, 
1976, p. 296). In 1976 an adult male was seen again at this locality on 
January 15-16 (J. J. Pujals) and March 6 (J. Greenberg) (Eisen- 
mann, Ridgely in litt.). The normal winter range is from Mexico to 
Nicaragua. | 


DENDROICA CERULEA (Wilson): Cerulean Warbler, Reinita Cerulea 


Sylvia cerulea Wilson, 1810, Amer. Ornith., 2, p. 141, pl. 17, fig. 5. (Philadelphia, 
Pennsylvania. ) 


Small; upper surface light blue (male) or bluish green (female) ; 
undersurface white or tinged yellow. 

Description.—Length 101-112 mm. Adult male, upper surface gray- 
ish blue, brighter on crown; a few fine black streaks on hindneck and 
back; rump light gray; upper tail coverts black, tipped light gray; les- 
ser wing coverts black, tipped blue; middle and greater wing coverts 
black, tipped white, forming two bars; remiges black, with outer web 
edged gray; rectrices black, with white on inner web extensive on outer 
pair, decreasing to a thin edge on innermost pair, and outer webs edged 
blue; black line from lores through eye; side of face light gray; under- 
surface white with narrow black band across upper breast and fine black 
streaks on sides. 

Adult female, crown bluish green; side of face and rest of upper 
surface light bluish olive-green, slightly lighter on rump; lesser wing 
coverts light green; middle and greater wing coverts black, tipped 
whitish, forming two bars; remiges black with outer web edged bluish 
olive-green and inner web edged white; rectrices as in male but blue 


276 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


on outer web replaced by light bluish olive-green; lores, superciliary, 
and entire undersurface white, extensively tinged pale dull yellow; 
underwing coverts white. 

Measurements.—Males (10 from the breeding range, taken in 
May), wing 62.5-69.5 (66.2), tail 40.5-45.5 (43.3), culmen from base 
10.3-12.4 (11.3), tarsus 14.8-16.7 (15.9) mm. 

Females (10 from the breeding range, taken in May), wing 61.0- 
65.0 (62.6), tail 39.9-44.1 (41.7), culmen from base 10.2-11.9 (11.2), 
tarsus 14.5-16.6 (15.8) mm. 

Migrant from the north. Uncommon. While this species winters in 
South America from Colombia and Venezuela south to Bolivia, and 
undoubtedly passes in part through Panama, I have seen only 7 speci- 
mens. The species is, however, hard to collect and passes through Pan- 
ama very quickly. The British Museum has 2 skins from Calovévora, 
Veraguas, taken by Arcé, and 3 others attributed to McLeannan, and 
therefore taken in the Canal Zone. One in the American Museum of 
Natural History is also from McLeannan. There is 1 in the Museum 
of Comparative Zoology collected on Isla del Rey (San Miguel), 
March 15, 1904, and 1 in the Smithsonian that Perrygo and I collected 
on Taboga Island, March 23, 1952. 

Ridgely (1976, p. 296) considers the Cerulean Warbler an “uncom- 
mon fall and rare spring transient (late August-early October; mid-to 
late March; one report from November 4) ... Only winter report is a 
sighting of a male on Cerro Campana on January 1, 1969 (Ridgely). 
... Usually arrives in small ‘waves’ in September when small groups 
may be encountered.” In early April 1976 Ridgely (1 litt.) recorded 
1 to 3 individuals per day at three different localities in the Canal Zone, 
with the maximum being 3 (2 males and a female) on the Pipeline Road 
on April 3. Extreme dates known to Eisenmann are August 20, 27, and 
April 3, 14. 


DENDROICA FUSCA (Miller): Blackburnian Warbler, 
Reinita Gargantianaranjada 


Motacilla fusca P.L.S. Miller, 1776, Natursyst., suppl., p. 175. (French Guiana.) 


Small; upper surface mainly black (grayish olive in female); throat 
and breast bright orange. 

Description.—Length 106-120 mm. Adult male, upper surface black, 
with small light orange patch in center of crown and two yellowish 
white stripes on back; wings black, with middle coverts broadly and 
greater coverts narrowly tipped white, forming a patch; primaries and 


FAMILY PARULIDAE Lag NG 


outer secondaries with yellowish white on edge of outer web, and in- 
nermost secondaries with white on edge of outer web; three outer pairs 
of rectrices white, tipped black, next pair black with white patch on in- 
ner web, remaining pairs black; lores, superciliary, side of neck, throat, 
and upper breast light orange, becoming bright orange in February; 
side of face and line through eye black; lower breast and rest of under- 
surface fading to pale orange-yellow; black stripes on side of breast 
and on sides. 

Adult female, upper surface grayish olive, streaked black, with small 
yellowish white patch in center of crown, often obscured; wings black, 
with middle and greater coverts tipped white, forming two bars; re- 
miges with outer web edged grayish olive; rectrices black with white 
patch on inner web of outer three pairs; lores and superciliary buff- 
yellow; side of face grayish olive; throat and upper breast from buff- 
yellow to light orange; rest of undersurface white, tinged buff, with a 
few black streaks on sides of breast and sides; underwing coverts 
white. 

Immature, like female, but paler. 

Measurements.—Males (10 from the breeding range, taken in 
May), wing 64.0-69.0 (67.4), tail 44.6-48.2 (46.4), culmen from base 
10.2-11.6 (11.1), tarsus 15.9-18.3 (17.4) mm. 

Females (10 from the breeding range, taken in May), wing 62.0- 
65.2 (63.8), tail 43.5-47.6 (45.7), culmen from base 10.7-12.2 (11.3), 
tarsus 16.4-18.3 (17.2) mm. 

Migrant and winter visitor from the north. Fairly common transient 
throughout on migration; a few winter in the highland forest of Chiri- 
qui. In some years it is more common as a winter resident in Chiriqut, 
equaling Black-throated Green Warblers in numbers. Asa migrant the 
Blackburnian Warbler has been recorded in fall from early September 
until November 17 (Eisenmann, Smiths. Misc. Coil., vol. 117, no. 5, 
1952, p. 51 and in spring from March 5, at Garachiné, Darién, in 1927 
(Griscom, Amer. Mus. Nov. no. 282, 1927, p. 10) to April 28, at Mt. 
Sapo, Darién, in 1941 (Bond and de Schauensee, Acad. Nat. Sci. Phila- 
delphia, Mon. no. 6, 1944, p. 42). I have also found this species at El 
Volcan and Santa Clara in Chiriqui during February. It has been re- 
corded as well from San José, Saboga, and Viveros in the Perlas 
Archipelago and at sea 96 km south of the Pearl Islands (specimen in 
the British Museum, taken October 24, 1924, by H. J. Kelsall). Some 
individuals may winter in the Darién highlands. D. fusca is usually 
seen in middle and high tree levels where it is often a part of mixed 
flocks of migrant warblers. 


278 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


A Blackburnian Warbler killed in a heavy rainstorm at Almirante, 
Bocas del Toro, on October 13, 1963, weighed 9.41 g and had an esti- 
mated 1.37 g of fat reserves, which might have enabled the bird to fly 
another 900 km (Rogers, Bird-Banding, 1965, pp. 115-116). 


[DENDROICA DOMINICA (Linnaeus): Yellow-throated Warbler, 
Reinita Gargantiamarilla 


Motacilla dominica Linnaeus, 1766, Syst. Nat., ed. 12, 1, p. 334. (Hispaniola.) 


Vagrant. This species is included on the basis of 3 sight reports: a 
well-marked male that I observed for several minutes on February 3, 
1958, at Almirante, Bocas del Toro, where it was feeding in a flowering 
tree with other warblers and orioles; a bird seen by C. Leahy, J. 
Gwynne, Ridgely, and Eisenmann on February 2, 1973, at Summit 
Gardens, Canal Zone (Ridgely, 1976, p. 296) (This bird was photo- 
graphed, but the subspecies was not determinable); and a bird seen by 
J. Baird and R. Forster on Cerro Azul, Province of Panama, on Janu- 
ary 28, 1978 (Ridgely, im litt.). The white lores and short bill character- 
istic of D. d. albilora were observed on the last-mentioned bird. D. a. 
albilora winters rarely south to Costa Rica.] 


DENDROICA PENSYLVANICA (Linnaeus): Chestnut-sided Warbler, 
Reinita de Lados Castanos 


Motacilla pensylvanica Linnaeus, Syst. Nat., ed. 12, vol. 1, 1766, p. 333. (Phila- 
delphia, Pennsylvania. ) 


Small; upper surface bright yellowish green; undersurface white, 
with, on male, chestnut stripe on side of breast and sides. 

Description.—Length 106-118 mm. Adult male, crown and back 
olive-yellow, with some black streaking on back, becoming more ex- 
tensive in March, when crown becomes bright yellow; rump black with 
feathers edged olive-yellow; upper tail coverts black, edged gray; wing 
coverts blackish, with lesser coverts tipped olive-yellow, middle and 
greater ones tipped white, tinged yellow, forming two bars; remiges 
blackish, edge of outer web of primaries gray, edge of outer web of 
secondaries olive-yellow; rectrices blackish with large white patch on 
inner web of outer two pairs; lores and side of head and neck gray un- 
til March, when lores, superciliary, and side of face in front of eye be- 
come black and rest of side of face becomes white; orbital ring white; 
undersurface white, with chestnut on sides becoming more extensive by 


FAMILY PARULIDAE 279 


March, when it is found on sides of breast also; underwing coverts 
white. 

Female and immatures, as male, but crown and back usually remain- 
ing olive-yellow; side of face remaining gray, and chestnut only on 
sides, often absent entirely. 

Mceasurements.—Males (10 from Panama), wing 61.2-66.5 (64.2), 
tail 45.7-50.6 (48.3), culmen from base 10.7-11.9 (11.2), tarsus 16.0- 
18.6 (17.3) mm. 

Females (10 from Panama), wing 56.1-61.0 (59.2), tail 43.6-48.8 
(46.0), culmen from base 10.5-11.3 (10.9), tarsus 15.8-18.0 (17.0) 
mm. 

Winter visitor from the north. Very common in the lowlands and 
less common in the highlands of Chiriqui; reported as common east 
through the mountains of Veraguas (Santa Fé, Chitra), ranging east 
through the Canal Zone to western San Blas (Mandinga) and eastern 
Province of Panama (Chico, Chepo). In the lowlands it is found 
widely, from Puerto Armuelles and Divala, Chiriqui; Paracoté, south- 
ern Veraguas; Almirante, Bocas del Toro; and Fl Uracillo, on the 
Caribbean slope of Coclé; to the Canal Zone. It is not reported to date 
from the eastern side of the Azuero Peninsula. I noted 2 on Isla Coiba 
January 12 and 29, 1956, and Ridgely (in litt.) saw 1 there on April 13, 
1976. E. A. Goldman wrote in his notes that in 1911 this was the ‘‘most 
abundant North American warbler in the northern half of the Canal 
Zone from January 12 until April 18 when the last specimen obtained 
was taken at Gatun.’ It decreases considerably after March. In 
Darién there are “few” records (Griscom, Bull. Mus. Comp. Zool., vol. 
73,1935, p. 368). 

The earliest record for Panama is September 7 (Willis and Eisen- 
mann, Smiths. Contrib. Zool., no. 291, 1979, p. 26); Ridgely (in litt.) 
has found individuals as late as April 25 (1980, Changuinola, Bocas 
del Toro) and May 1 (1976, Ft. Sherman, Canal Zone). Evidently 
many individuals remain in the same locality for several months—of 
those banded by Loftin et al. (Bird-Banding, 1966, p. 41) at Curundu, 
in the Canal Zone, 1 banded December 8, 1963, was taken at the same 
locality February 9, 1964. Another banded at Curundu December 3, 
1963, was recaptured there April 2, 1966 (Loftin et al., Bird-Banding, 
MGA pebot):, 

The Chestnut-sided Warbler is found most often in second-growth 
woodland, clearings, and edges, often in groups with other migrant 
warblers. 


280 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


In a banding study at Almirante, Rogers and Odum (Auk, 1966, p. 
418) found that Chestnut-sided Warblers arriving there in fall had 
exhausted their migratory fat reserves and had lost some weight from 
the breast muscles. The fat-free weight of 20 was 6.8-9.1 (8.0) grams. 
One collected by Strauch (Bull. Brit. Orn. Club, 1977, p. 64) weighed 
102 

Chestnut-sided Warblers (fide E. S. Morton) molt much ahead of 
Bay-breasted Warblers, with which they often associate in Panama, 
and by mid- or late March are already in full breeding plumage. They 
are often seen with other passerines feeding on the fruit of the abun- 
dant Miconia argentea. Usually a mixed feeding aggregation of paru- 
lids will include only 1 Chestnut-sided Warbler. 

Eisenmann reports that birds seen in Panama usually carry the tail 
diagonally cocked above the back, and sometimes jerk it nervously. 
The wings are often carried drooped below the body. 


DENDROICA CASTANEA (Wilson): Bay-breasted Warbler, 
Reinita Pechicastana 


Sylvia castanea Wilson, Amer. Orn., vol. 2, 1810, p. 97, pl. 14, fig. 4. (Pennsyl- 
vania. ) 

Small; upper surface green (grayish with brown crown in males 
after February); undersurface buff, sometimes with brown on throat 
and sides. 

Description —Length 111-130 mm. Male, September through Feb- 
ruary, crown and upper back light yellowish olive, finely streaked with 
black; lower back, rump, and upper tail coverts gray; wings black, with 
lesser coverts tipped gray, middle and greater coverts tipped white, 
forming two bars, remiges with inner web edged white, outer web of 
primaries 9-4 edged gray, outer web of rest of remiges light yellowish 
olive; rectrices black with large white spot on inner web of outer two 
pairs; lores and orbital ring white; side of head light yellowish olive; 
undersurface white, tinged buff, most heavily on abdomen and under- 
tail coverts; flanks sometimes russet; underwing coverts white. 

Male, March and April, forecrown, side of head, including area all 
around eye, and chin, black; hindcrown and nape russet; upper back 
streaked black and warm buff; side of neck warm buff; throat, breast, 
sides, and flanks russet; rest of undersurface buffy white; lower back, 
rump, wings, and tail as above. 

Iemale, like September-February male; in March and April some- 
times with russet on crown, throat, and breast. 


Mcasurements.—Males (10 from Panama), wing 69.9-77.0 (73.7), 


FAMILY PARULIDAE 281 


tail 48.0-55.1 (52.2), culmen from base 11.1-12.1 (11.6), tarsus 17.7- 
LORElS:Z) mm. 

Females (10 from Panama), wing 65.0-75.1 (69.7), tail 47.0-53.8 
(50.0), culmen from base 10.3-12.8 (11.6), tarsus 16.0-18.7 (17.1) 
mm. 

Migrant and winter visitor from the north. Common. Found on the 
Pacific slope from eastern Veraguas, the mountain area of Cocle (El 
Valle), and western Province of Panama (Cerro Campana) eastward, 
being especially common in the Bayano River Valley of eastern Pan- 
ama Province and in Darién; it has not been recorded from Chiriquti. 
On the Caribbean side it ranges throughout the Republic from western 
Bocas del Toro to eastern San Blas. Mainly found in the Tropical 
Zone, it is recorded in the lower edge of the Subtropical Zone at El 
Valle, and on Cerro Campana. In the Perlas group it has been recorded 
from the islands of San José, Isla del Rey, and Saboga. 

The first Bay-breasted Warblers arrive in Panama in mid-September 
(Ridgely, 1976, p. 297), although few are seen before October. Molt 
of adults takes place in March; immature birds seem to molt slightly 
later. By April, numbers of Bay-breasted Warblers have decreased in 
Panama. The latest date for which the species is recorded in Panama 
is May 1, when in 1969 G. Tudor saw 1 at Cerro Azul, Province of 
Panama. On March 18 and 19, 1957, I found a considerable migration 
through the forest along the lower Rio Oria west of the southeastern 
tip of the Azuero Peninsula. While D. Hicks netted this species almost 
daily at Almirante, Bocas del Toro, during October 15-26, 1964, he 
caught none there on the following spring migration (March through 
April 1965), suggesting different routes on the southward and north- 
ward flights (Eisenmann im litt.). N.G. Smith reports seeing one at 
Almirante during the “last week of March” 1965. 

The Bay-breasted Warbler is the commonest wintering warbler in 
the forests of central Panama, especially on the Caribbean slope. Its 
numbers are often underestimated, as the birds are usually in forest 
canopy or high in woodland and easily overlooked. Central and eastern 
Panama are the centers of its notably small wintering range. Its popu- 
lation may have increased in the 1970’s in response to spruce bud-worm 
outbreaks in its breeding area. In Panama it feeds largely on the fruit 
of Micomia argentea and of Bursera, two common small trees. Ridgely 
(in litt.) estimated 60 Bay-breasted Warblers on the Achiote Road, 
Canal Zone, March 18, 1980, and 50 at Ft. Sherman the next day. 

A male collected by Strauch (Bull. Brit. Orn. Club, 1977, p. 64) in 
Province of Panama (location not given) weighed 11.4 g. 


282 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


DENDROICA STRIATA (Forster): Blackpoll Warbler, Reinita Coroninegra 
Muscicapa striata J.R. Forster, 1772, Phil. Trans. vol. 62, art. 29, read June 18 
and 25), pp. 406, 428. (Fort Severn, Ontario.) 

Small; both sexes in basic plumage, dull yellowish green streaked 
with black on upper surface; undersurface white, tinged yellow, some- 
times slightly streaked with black. Adult male in alternate plumage, 
streaked black and white. 

Description —Length 115-126 mm. Adult male in basic plumage, 
side of head and upper surface olive-green, becoming gray on rump 
and upper tail coverts, and finely streaked with black, especially on 
crown and upper back; wing coverts blackish, with middle and greater 
ones tipped white, forming two wing-bars; remiges blackish, with 
outer webs edged olive-green; tertials edged whitish; rectrices blackish 
with white patches on webs of outer two pairs; undersurface white, 
tinged yellow on sides of throat, across breast, and on sides and flanks, 
and finely streaked with black on sides and flanks; underwing coverts 
white. 

Adult male in alternate plumage, crown black; nuchal band streaked 
black and white, rest of upper surface light gray, becoming brownish on 
rump and upper tail coverts; upper back streaked with black; wings and 
tail as in basic plumage; side of head and undersurface white, with a 
broad black streak running from chin, along side of throat, to side of 
breast; sides and flanks streaked with black. 

Adult female in basic plumage, like male in basic plumage, but with 
less streaking on back, often absent from crown, and yellow tinge more 
extensive on undersurface. 

Adult female in alternate plumage, like male in basic plumage. 

Measurements.—Males (10 from North America, taken in May), 
wing 69.5-75.5 (72.5), tail 48.5-52.7 (50.1), culmen from base 11.1- 
12.6 (12.1), tarsus 17.1-19.6 (18.8) mm. 

Females (10 from North America, taken in May), wing 68.0-73.0 
(69.5), tail 47.3-50.7 (48.1), culmen from base 11.5-12.4 (12.0), tar- 
sus 18.2-19.6 (18.9) mm. 

Accidental. There is an adult male specimen in full spring plumage 
in the British Museum marked as taken in Panama by McLeannan 
that was received in the Tweedale Collection with no further data. The 
form of the skin, in the arrangement of wings and feet and the manner 
in which the bill is closed, is typical of the method employed by Mc- 
Leannan, so that there is no reason to doubt the locality. This collector’s 
activities centered along the line of the Panama Railroad, mainly on 
the Atlantic slope near the former station of Lion Hill (now covered 


FAMILY PARULIDAE 283 


by Gattin Lake). The record, therefore, is to be attributed to the Canal 
Zone (see Wetmore, Auk, 1958, pp. 467-468). This species migrates 
through the West Indies and is only of accidental or, at most, casual 
occurrence in Middle America. 

Eisenmann (Smiths. Misc. Coll., vol. 117, no. 5, 1952, p. 51) de- 
scribes 2 sight reports for this species on Barro Colorado Island: “Dr. 
A. A. Allen writes that November 6, 1944, he observed one ‘close 
enough so that I had no trouble seeing the light-colored legs and white 
under tail coverts.. Mrs. G. G. Fry writes that on February 8 and 9, 
_ 1950 she heard a song which she recognized as that of this species, and 
that one of the members of her party, Mrs. L. J. Francke, saw the bird 
and said it was a male in spring plumage.” Ridgely comments, how- 
ever, that some autumn Bay-breasted Warblers have the characters 
Allen noted and that in February Blackpoll Warblers are not in “spring 
plumage”’ or likely to be singing. On October 19 and 29, 1964, indi- 
viduals identified as this species by D. L. Hicks were banded and re- 
leased at Almirante, Bocas del Toro (Ridgely, 1976, p. 297). There 
are 2 recent sightings from Costa Rica, in December 1977 and January 
1975 (Stiles and Smith, Brenezia, vol. 17, 1980, p. 151), and a speci- 
men was collected there by F. G. Stiles in “fall” of 1980 (in litt. to 
Eisenmann). 


DENDROICA PALMARUM PALMARUM (Gmelin): Palm Warbler, 
Reinita del Palmar 


Motacilla palmarum Gmelin, 1789, Syst. Nat., 1(2), p. 951. (Santo Domingo.) 


Small; upper surface brown; undersurface buffy with fine brown 
streaks, yellow on undertail coverts and sometimes on throat. 

Description.—Length 109-126 mm. Adult (sexes alike), crown and 
back olive-brown, finely streaked with darker brown; forecrown be- 
coming russet by March; rump and upper tail coverts yellow; wing 
coverts blackish brown, tipped buffy brown; remiges blackish brown, 
with outer web of primaries narrowly edged yellow, and outer edges of 
secondaries broadly edged buffy brown; rectrices blackish brown with 
large white patch on inner web of outermost pair and small white patch 
on next pair; superciliary whitish, becoming yellow by March; orbital 
ring whitish; undersurface whitish or olive-buff, finely streaked with 
brown on breast and sides of throat; undertail coverts lemon yellow; 
throat becomes lemon yellow by March, rest of undersurface sometimes 
tinged yellow as well. 

Measurements—Males (10 from the breeding range, taken in May 


284 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


and July), wing 61.0-67.1 (64.4), tail 46.2-53.2 (49.9), culmen from 
base 10.3-11.9 (11.3), tarsus 18.3-19.6 (19.1) mm. 

Females (10 from the breeding range, taken in May and July), wing 
59.4-62.1 (61.0), tail 46.1-49.3 (47.6), culmen from base 10.8-11.8 
(11.2)¢ tarsus 18.3219.7 \ (1911 ),mmt 

Winter visitor from the north. Rare. This species was not known 
from Panama before 1965. The normal winter range is south to Hon- 
duras and northeastern Nicaragua. It is found more rarely to Costa 
Rica. On March 15, 1965, a male was collected by Dr. A. Diaz near 
Tocumen, on the Pacific slope about 32 km east of Panama City, Proy- 
ince of Panama. The specimen is now in the Smithsonian. Hicks et ai. 
(Condor, 1967, p. 319) erroneously list the date as October 14. Other 
sight records led Ridgely (1976, p. 297) to describe the Palm Warbler 
as “a rare but probably regular winter visitor in small numbers on 
Caribbean slope of Canal Zone and savannas of eastern Panama Proy- 
ince . . . mid-November-mid-March . . . usually on or very near the 
ground. In Canal Zone, prefers lawns in residential areas.” 


[DENDROICA DISCOLOR (Vieillot ): Prairie Warbler, 
Reinita de la Pradera 


Sylvia discolor Vieillot, 1808 or 1809, Hist. Nat. Ois. Amer. Sept., 2 (1807), p. 37, 
pl. 98. (United States and Greater Antilles.) 


This species is included on the basis of a sight report of 1 at Volcan, 
Chiriqui on January 23, 1973, found by Richard Brownstein and Wal- 
ter George. Brownstein (in litt. to Wetmore, June 25, 1974) informs 
me that they observed the bird for 10 minutes as close as 7 m; they did 
not, however, supply any details of plumage. The Prairie Warbler 
normally winters in Florida, the West Indies, and, in very small num- 
bers, from Yucatan to Nicaragua, mainly on islands off the Caribbean 
coast, and on islands off the north coast of South America. There are 
also some recent sight reports from Costa Rica (Stiles and Smith, 
Brenezia, vol. 17, 1980, p. 15).] 


SEIURUS AUROCAPILLUS (Linnaeus): Ovenbird, Reinita Suelera 


Medium size; upper surface mostly light yellowish olive with orange- 
brown central crown stripe; undersurface white, streaked with black. 

Description.—Length 130-138 mm. Adult (sexes alike), side of head 
and entire upper surface, including wing coverts, light yellowish olive, 
with broad central crown stripe of orange-brown, bordered by narrow 


FAMILY PARULIDAE 285 


blackish stripes; remiges dusky, edged light yellowish olive; rectrices 
light yellowish olive; eye-ring white; undersurface white with narrow 
blackish streak on side of throat and blackish streaks across breast and 
down sides and flanks; underwing coverts white, tinged yellow. 

The North American Ovenbird is a fairly common migrant and 
winter resident in western Panama, becoming uncommon to rare to the 
east. In Darién it has been recorded only twice (Griscom, Bull. Mus. 
Comp. Zool., vol. 78, 1935, p. 368). I have always found it in dry for- 
est, but it has been collected from sea level to 2900 m, on Volcan de 
Chiriqui (Bangs, Proc. New England Zool. Club, vol. 3, 1902, p. 62). 

The Ovenbird has been found in Panama from late September until 
April 25 (Loftin and Olson, Carib. Journ. Sci., vol. 3, no. 4, 1963, p. 
195). There are numerous recapture records from the Gorgas Memo- 
rial Laboratory banding station at Almirante, Bocas del Toro. A par- 
ticularly notable one is of a bird banded there October 13, 1963, and re- 
captured April 23,1965; and again on October 17, 27, and 28, 1965 
(Loftin et al., Bird-Banding, 1967, p. 151). Another study done at 
Almirante found the weight of 87 fall migrant Ovenbirds to range from 
12.9-19.0 (15.74) g and that the birds had exhausted their migratory 
fat reserves (Rogers and Odum, Wilson Bull., 1966, p. 418). Willis 
(Living Bird, 1966, p. 206) noted that on Barro Colorado Island, 
Canal Zone, where the Ovenbird is rare, it normally ignores army ants. 
There it has been recorded from October 18 to March 31 (Willis and 
Eisenmann, Smiths. Contrib. Zool. no. 291, 1979, p. 26). 

Two subspecies have been identified as occuring in Panama, auro- 
capillus and furvior. 


SEIURUS AUROCAPILLUS AUROCAPILLUS (Linnaeus) 
Motacilla aurocapilla Linneaeus, 1766, Syst. Nat., ed. 2, 1, p. 334. (Pennsylvania.) 


Seiurus aurocapillus canivirens Burleigh and Duvall, 1952, Wilson Bull., 64, p. 39. 
(Margaret, Fannin County, Georgia.) 


Characters.—Upper surface more greenish; central crown stripe 
more yellowish. 

Measurements.—Males (10 from the breeding range, taken in May), 
wing 73.3-80.6 (78.0), tail 52.7-58.5 (56.7), culmen from base 13.2- 
15.0 (13.8), tarsus 19.3-23.7 (20.9) mm. 

Females (10 from the breeding range, taken in May), wing 69.5- 
77.2 (73.4), tail 51.3-57.6 (53.9), culmen from base 12.7-14.7 (13.5), 
tarsus 19.3-22.1 (20.9) mm. 

Migrant and winter visitor from the north. The Smithsonian has 7 


286 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Panamanian specimens of this race: 4 were collected in the Chiriqui 
highlands, 1 at Almirante, Bocas del Toro, and 1 each from Isla Cébaco 
and Isla Gobernadora in the Golfo de Montijo off the Pacific Coast of 
Veraguas. 


SEIURUS AUROCAPILLUS FURVIOR Batchelder 


Seiurus aurocapillus furvior Batchelder, 1918, Proc. New England Zool. Club, 6, 
p. 81. (Near Deer Pond, Newfoundland.) 


Characters.—Upper surface browner than in nominate race; streak- 
ing on undersurface heavier and blacker; flanks darker. 

Measurements.—Males (10 from the breeding range, taken in May, 
June, and July), wing 76.5-82.0 (79.0), tail 54.6-58.1 (56.7), culmen 
from base 12.5-15.5 (14.0), tarsus 20.9-23.2 (21.8) mm. 

No specimens of females were available for measurement. 

Migrant and winter visitor from the north. This very weakly dif- 
ferentiated race breeds exclusively in Newfoundland, Canada. Pan- 
ama appears to be the southern limit of its winter range. At least 2 
specimens in the collections of the Smithsonian from Panama have been 
identified as furvior: 1 taken October 18, 1962, at Almirante, Bocas 
del Toro, and the other taken March 10, 1952, at Chilar, Colon. 


SEIURUS NOVEBORACENSIS (Gmelin): Northern Waterthrush, 
Reinita Arrollera del Norte 


Motacilla noveboracensis Gmelin, 1789, Syst. Nat., 1(2), p. 958. (Louisiana and 
New York.) 


Small; upper surface olive; undersurface yellowish, streaked with 
olive. 

Description.—Length 121-138 mm. Adult (sexes alike), upper sur- 
face, including wings and tail, olive; superciliary, broken eye-ring, and 
sometimes stripe on forehead, buffy white; side of face and stripe 
through eye olive; throat white, rest of undersurface buffy yellow, 
streaked olive except on center of abdomen; underwing coverts light 
olive-brown. 

Measurements.—Males (10 from Panama), wing 68.8-77.9 (74.8), 
tail 46.8-54.7 (51.3), culmen from base 12.0-14.1 (13.2), tarsus 19.4- 
Awe Alli \raacnay, 

Females (10 from Panama), wing 69.5-75.0 (71.6), tail 45.1-53.1 
(49.4), culmen from base 12.0-13.9 (13.1), tarsus 19.8-22.1 (20.7) 
mm. 

Migrant and winter visitor from the north. Very common through- 


FAMILY PARULIDAE 287 


out in the lowlands of both slopes, particularly in mangroves and near 
water, both running and still. I have also seen them among the jumbled 
rocks along the beach on San José in the Pearl Islands, where they are 
common throughout the group, and on the ocean beach at Isla Goberna- 
dora, Veraguas. In migration they are encountered away from water 
as well, on the paths in dry woods, and in the highlands. On Volcan 
de Chiriqui, Monniche (Blake, Fieldiana: Zool., vol. 36, no. 5, 1958, 
p. 559) collected Northern Waterthrushes as high as 1590 m during 
October, and W. W. Brown, Jr. (Bangs, Proc. England Zool., Club, 
vol. 3, 1902, p. 62) collected a male at Boquete (1200 m) on March 27, 
1901. Eisenmann saw 1 above Cerro Punta, Chiriqui at approximately 
2100 m on September 19, 1958. Northern Waterthrushes arrive in 
Panama in late September and most depart by late April. Extreme 
dates known to Fisenmann (im litt.) are September 11 to May 13 (both 
Ft. San Lorenzo, Canal Zone). In the exceptionally long dry season of 
1977, however, N. G. Smith saw 2 together at Summit Gardens on May 
26 and June 2; several other northern migrants also remained late. At 
Almirante, Bocas del Toro, in the southward migration of 1964, 15 were 
netted on September 29 and 16 on October 10; the following spring 
numbers netted were much fewer, not exceeding 2 or 3 in any one day 
between March 21 and April 23 (D. L. Hicks in htt. to Eisenmann). 

Banding results suggest that many of the individuals that do not con- 
tinue on to South America are highly sedentary while in Panama; 
Loftin et al. (Bird-Banding, 1966, p. 42) banded a Northern Water- 
thrush in the Canal Zone October 5, 1963, that was recaptured there on 
Iebruary 9, 1964. One banded at Almirante, November 10, 1962 was 
retaken there January 7, 1963, and again on October 14, 1963, indicat- 
ing that some individuals return to the same wintering area in subse- 
quent years, as they have been shown to do in Venezuela (Schwartz, 
Living Bird, 1964, pp. 169-184). Of the 65 birds banded by Loftin 
and his associates in the Canal Zone and the 338 banded at Almirante 
in 1963 and 1964, there were 5 repeats in the Canal Zone and 15 at 
Almirante. In another banding study at Almirante, Rogers and Odum 
(Wils. Bull., 1966, p. 418) weighed 165 Northern Waterthrushes, 
which ranged between 10.5-19.2 (14.69) g, and found that, unlike sev- 
eral other migrant species, including the Ovenbird, the waterthrushes 
had not exhausted their fat reserves. 

Willis (Living Bird, 1966, p. 207) observed individuals picking at 
insects flushed by swarms of army ants on Barro Colorado Island; 
however, the birds were never persistent swarm followers. The weights 
of 7 collected by Burton (Bull. Brit. Orn. Club, 1975, p. 85) ranged 


288 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


from 13.2 to 16.0 g. A female taken at Fort Sherman, Canal Zone, by 
G. V. N. Powell (in litt. to Eisenmann) weighed 18 g. 


SEIURUS MOTACILLA (Vieillot): Louisiana Waterthrush, 
Reinita Arrollera de Louisiana 


Turdus motacilla Vieillot, 1808, Hist. Nat. Ois. Amer. Sept., 2 (1807), p. 9, p. 65. 
(Kentucky. ) 


Small; upper surface olive-brown; undersurface white, streaked 
olive-brown across breast and sides. 

Description.—Length 129-141 mm. Adult (sexes alike), upper sur- 
face, including wings and tail, line through eye, and side of face, olive- 
brown; superciliary white, extending to nape; line around lower por- 
tion of eye, moustachial stripe, and undersurface white, becoming buffy 
on abdomen and undertail coverts, and with olive-brown streaks on 
breast and sides; underwing coverts buffy brown. 

Measurements.—Males (9 from Panama, Costa Rica, and Colom- 
bia), wing 80.1-83.4 (82.2), tail 44.6-54.4 (50.4), culmen from base 
14.4-16.9 (15.5), tarsus 21.5-22.7 (22.2) mm. 

Females (9 from Panama), wing 72.5-82.2 (77.5), tail 46.7-54.7 
(50.0), culmen from base 14.2-16.4 (15.5, average of 8), tarsus 21.3- 
23.1 (22.4) mm. 

Migrant and winter visitor from the north. Uncommon as a migrant, 
chiefly in mid-August to September and in March, when it may be found 
throughout the Republic, and rather rare as a winter resident, mostly 
in highlands of Chiriqui. The Louisiana Waterthrush prefers running 
water. The first to arrive in Panama appear by early August—long be- 
fore any Northern Waterthrushes—and some linger into early April 
(Ridgely, 1976, p. 298). Most have left Panama by mid-March; later 
sight reports may be misidentified Northern Waterthrushes. The lat- 
est known report from Panama is a specimen netted at Almirante, 
Bocas del Toro, April’ 3, 1965)() Li) Hicks) Unity” Georetaicollae 
Monniche (Blake, Fieldiana: Zool., vol. 36, no. 5, 1958, p. 559) col- 
lected a female at 1650 m on the Volcan de Chiriqui. On the Pearl 
Islands W. W. Brown, Jr., (Thayer and Bangs, Bull. Mus. Comp. 
Zool., vol. 46, 1905, p. 156) collected 1 on San Miguel on March 18, 
1904, and I saw 1 on San José on March 7, 1946. 

If the absence of fat can be taken as an indication that a bird is not 
yet migrating, then individuals I collected at the Peluca Hydrographic 
Station in the Province of Panama on February 21, 1961, and on the 
south fork of the Rio Pucro on Cerro Mali, Darién, on March 2, 1964, 
were winter residents. On Barro Colorado Island, Willis (Living Bird, 


FAMILY PARULIDAE 289 


1966, pp. 206-207) found this species an “uncommon winter resident” 
that ignored swarms of army ants when they passed by. 


OPORORNIS FORMOSUS (Wilson): Kentucky Warbler, 
Reinita Hermosa 


Sylvia formosa Wilson, 1811, Amer. Ornith., 3, p. 85, pl. 25, fig. 3. (Kentucky.) 


Small; upper surface yellowish olive, with black markings on head; 
undersurface bright yellow. 

Description—Length 114-126 mm. Adult male, forecrown black, 
with feathers often tipped gray; line from bill to eyebrow and continu- 
ing around rear of eye to lower rim, yellow; lores, side of face, and 
side of throat black; rest of upper surface, including tail, yellowish 
olive; primaries dusky, with outer webs edged yellowish olive; rest of 
wing yellowish olive; undersurface yellow, with yellowish olive some- 
times extending from sides to flanks; bend of wing and underwing 
coverts yellow. 

Adult female, as male, but black areas of head less extensive, often 
replaced by dusky brown. 

Measurements.—Males (10 from Panama), wing 66.0-71.5 (68.7), 
tail 45.3-53.3 (48.7), culmen from base 10.4-13.3 (12.0), tarsus 19.4- 
227 (214) mm. 

Females (9 from Panama, Costa Rica, and Colombia), wing 64.0- 
69.6 (65.9), tail 43.3-48.7 (47.2), culmen from base 11.6-13.2 (12.3), 
tarsus 19.9-22.9 (21.8) mm. | 

Migrant and winter visitor from the north. Fairly common, but in- 
conspicuous, throughout in forest and second-growth woodland, where 
it inhabits thickets of shrubbery, similar to its northern breeding habi- 
tat. Although during migration it occurs in the highlands, where it has 
been collected by W. W. Brown, Jr., as high as 2310 m, near Boquete, 
Chiriqui, on February 22, 1901 (Bangs, Proc. New England Zool. Club, 
vol. 3, 1902, p. 62), most wintering individuals are found in the low- 
lands (Ridgely, 1976, p. 299). The Kentucky Warbler has been re- 
corded in Panama from September 4 (Willis, Living Bird, 1966, p. 
207) to April 28, (Loftin and Olson, Carib. Journ. Sci., vol. 3, no. 4, 
1968, p: 195... 

Banding results indicate that individuals wintering in Panama are 
highly sedentary and faithful to a site, returning in subsequent years. 
One banded at Almirante, Bocas del Toro, on October 14, 1963, was 
retaken there February 25, 1964, and another banded there October 12, 
1963, was taken again October 29, 1964 (Loftin et al., Bird-Banding, 
1966, p. 42). In the Canal Zone, a Kentucky Warbler banded near the 


290 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Gamboa horse stables October 22, 1964, was retaken there March 5, 
1965, and 1 banded on the Pipeline Road near Gamboa November 7, 
1964, was recaptured there October 21, 1965. One banded by Karr 
(Bird-Banding, 1971, p. 299) on the Chiva Chiva Road, Canal Zone, 
on December 14, 1968, was recaptured 100 m from the same spot on 
January 27 and 28, 1971. 

Willis (Living Bird, 1966, pp. 207-208) found that Kentucky War- 
blers were regular followers of army ant swarms on Barro Colorado 
Island. They usually stay ahead or to the side of the ants, but often as- 
sociate with them for more than 5 minutes at a stretch. If 2 individuals 
are near a single group of ants they usually stay at different ends of the 
swarm. When not searching for insects flushed by ants, Kentucky 
Warblers often associate with or follow Spotted Antbirds (Hylophylax 
naevioides) or with wandering mixed flocks. On March 15, 1955, at 
FE] Volcan, Chiriqui I saw 1 with a moving flock of warblers; prob- 
ably all were migrants. One collected by Burton (Bull. Brit. Orn. Club, 
1975, p. 85) weighed 13.5 g. Ridgely (1976, p. 299) points out that 
this species and the Ovenbird are the only 2 North American migrant 
warblers that regularly inhabit the interior of mature forests in winter; 
they are, therefore, more sensitive to habitat destruction than most other 
wintering species, which usually inhabit more or less disturbed areas 
when in Panama. 

Raikow’s (Bull. Carnegie Mus. Nat. Hist. no. 7, 1978, p. 35) myo- 
logical studies have confirmed the traditional separation of the genus 
Oporornis from Geothly pis, into which the former had been placed by 
Lowery and Monroe (Check-list Birds World, vol. 14, 1968, p. 38). 


OPORORNIS AGILIS (Wilson ): Connecticut Warbler, 
Reinita de Connecticut 


Sylvia agilis Wilson, 1812, Amer. Ornith., 5, p. 64, pl. 39, fig. 4. (Connecticut. ) 


Small; upper surface yellowish olive; head and breast gray; under- 
surface yellow. 

Description.—Length 118-131 mm. Adult male, crown gray, with 
feathers often tipped yellowish olive; rest of upper surface, including 
tail, yellowish olive, slightly lighter on rump and upper tail coverts; 
primaries dusky, with outer web edged yellowish olive; rest of wing 
yellowish olive; complete orbital ring white; throat, upper breast, and 
sides of head gray, slightly lighter on throat; sides and flanks light yel- 
lowish olive; rest of undersurface yellow; bend of wing yellow; under- 
wing coverts light yellowish olive. 


FAMILY PARULIDAE 291 


Female and immatures, as adult male, but gray replaced by drab 
grayish brown, slightly lighter on throat; orbital ring whitish to yel- 
lowish. 

Measurements.—Males (10 from North America, taken in May), 
wing 69.5-74.0 (71.8), tail 45.8-50.9 (48.1), culmen from base 10.6- 
12.9 (11.9), tarsus 19.6-21.4 (20.9) mm. 

Females (8 from North America, taken in May and June), wing 
67.0-71.0 (68.5), tail 46.2-49.4 (48.2), culmen from base 11.3-12.3 
(11.6), tarsus 18.7-21.5 (20.1) mm. 

Migrant from the north. Very rare, apparently only a transient; 
recorded only from the lowlands of western Bocas del Toro, between 
late September-late October and late March-early April. The occur- 
rence of this species in Panama has only been confirmed recently, when 
3 specimens taken in mist nets were collected at Almirante. A female 
was taken on October 15, 1963, and an unsexed individual on Septem- 
ber 30, 1964 (Hicks, Méndez, and Loftin, Condor, 1967, p. 319). The 
third, a female with the skull ossified, was collected by T. V. Heatley 
on October 23, 1967; (AMNH no. 801541). The spring reports are 
based only on sightings (Ridgely, 1976, p. 299). The Connecticut 
Warbler winters in Venezuela, eastern Colombia, and northern Brazil; 
the bulk of the population travels to and from North America via the 
West Indies or across the Caribbean. D. L. Hicks wrote Eisenmann 
that, in addition to the bird he preserved at Almirante in September 
1964, he banded and released other individuals identified as this species 
on September 28 and 29 and October 3, and in the following spring on 
March 21, 22, 28, and 29, and April 4, 5, and 9, 1965. 


OPORORNIS PHILADELPHIA (Wilson): Mourning Warbler, 
Reinita Encopetada 


Sylvia Philadelphia Wilson, 1810, Amer. Ornith., 2, p. 101, pl. 14, fig. 6. (Within a 
few miles of Philadelphia, Pennsylvania. ) 


Small; head gray, male with black on breast; rest of upper surface 
yellowish olive; undersurface yellow. 

Description.—Length 111-120 mm. Adult male, entire head slate 
gray; rest of upper surface, including tail, yellowish olive; primaries 
dusky with outer webs edged yellowish olive; rest of wing yellowish 
olive; center of throat and breast black, with feathers edged gray; 
sides of throat and breast gray; sides and flanks light yellowish olive; 
rest of undersurface, edges of wing, and underwing coverts yellow. 

Adult female, like male, but head deep grayish olive and throat and 
upper breast drab. 


292 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


Immatures, upper surface buffy olive; undersurface yellow, tinged 
brown on throat and upper breast. 

Measurements.—Males (10 from Panama), wing 58.9-65.0 (61.5), 
tail 45.9-50.7 (48.6), culmen from base 10.2-12.7 (11.4), tarsus 17.8- 
Ze 2 CLO 38) saan 

Females (8 from Panama), wing 55.4-60.1 (57.5), tail 45.5-51.7 
(46.7), culmen from base 10.7-12.2 (11.4), tarsus 19.3-21.1 (20.4) 
mm. 

Migrant and winter visitor from the north. Fairly common through- 
out as a transient, less common as a winter resident. The Mourning 
Warbler is found in Panama from mid-September to mid-May: F. S. 
Blanton collected a male at Gamboa in the Canal Zone on September 
16, 1953, and Monniche (Blake, Fieldiana: Zool., vol. 36, no. 5, 1958, 
p- 559) collected a male at 1590 m on the Volcan de Chiriqui on May 
16. E. A. Goldman’s notes remark that this species was “rather com- 
mon in the northern half of the Zone between January 11 and April 24 
[1911] when the last one seen was shot at Gatun. The disappearance 
of the migrating warblers and some other North American birds 
seemed to be closely coincident with the opening of the rainy season 
about April 24, as indicated by a thunder shower and change in the pre- 
vailing direction of the wind. The wind about this time began to come 
from southward or southwestward and the northerly trade ceased to 
blow.” Eisenmann saw 3 at Ft. Davis, Canal Zone, as late as May 11, 
1960. 

I have encountered this species throughout the Republic, at El Vol- 
can, Santa Clara, Concepcion, Buena Vista, and Alanje in Chiriqui; 
Almirante, Bocas del Toro; El Valle and El Uracillo, Coclé; Chilar in 
Colon; several localities in the Canal Zone and eastern Province of 
Panama; on the Rio Jaqué in Darién; and Mandinga and Puerto 
Obaldia in San Blas. Ridgely (1m litt.) finds that during the northern 
winter it is numerous in the highlands of Chiriqui and Darién. It is 
always in dense shrubbery or in an area of weeds bordering taller 
growth. 

Banding studies indicate that Mourning Warblers wintering in Pan- 
ama are faithful to a site through the winter and in succeeding years. 
Loftin et al. (Bird-Banding, 1966, p. 42) had the following returns at 
Almirante: April 19, 1963, retaken February 25, 1964; October 9, 1963, 
retaken September 27, 1964; and November 1, 1963, retaken April 21, 
1964. One banded at Gamboa January 27, 1964, was retaken there 
November 11, 1964. Five collected by Burton (Bull. Brit. Orn. Club, 
1975, p. 86) at El Real, Darién weighed from 9.5 to 10.0 g. G. V.N. 


FAMILY PARULIDAE 293 


Powell (1m litt. to Eisenmann) took a female immature (skull unossi- 
fied) on October 19, 1966, at It. Sherman that weighed 11 g. 


OPORORNIS TOLMIEI TOLMIEI (Townsend): MacGillivray’s 
Warbler, Reinita de Tupidero 


Sylvia Tolmiei J.K. Townsend, 1839 (Apr.), Narr. Journey Rocky Mountains, 
etc. p. 343. (Fort Vancouver, Clarke County, Washington.) 


Small; head, throat, and chest slate gray; upper surface yellowish 
olive; undersurface yellow. 

Description—Length 114-127 mm. Adult male, crown and nape 
slate gray; rest of upper surface, including tail, yellowish olive; pri- 
maries dusky, with outer web edged yellowish olive; rest of wing yel- 
lowish olive; lores black; broken eye-ring white; sides of throat and 
upper breast light gray; center of throat and chest black, with most 
feathers tipped white; sides and flanks light yellowish olive; rest of 
undersurface, bend of wing and underwing coverts yellow. 

Adult female, like male, but crown grayish olive; throat whitish, 
tinged gray; upper breast light grayish olive. 

Measurements.—Males (10 from the breeding range, taken in May 
and June), wing 57.0-61.9 (59.4), tail 50.0-53.5 (51.9), culmen from 
bases t@(5-11.2 (10:8), tarsus. 18:2-21.2 (20.1). mm: 

Females (10 from the breeding range, taken in June and July), 
wing 55.0-61.0 (56.4), tail 46.5-52.4 (49.5), culmen from base 10.6- 
Ori 2) tarsus 18.8-21 1 (19.9)) mm. 

Winter visitor from the north. Uncommon, recorded only from 
Chiriqui and the Canal Zone. In Chiriqui, the MacGillivray’s Warbler 
is known mainly from the highlands, where Monniche collected it at 
the Volcan de Chiriqui between 1590 and 1800 m (Blake, Fieldiana: 
Zool., vol. 36, no. 5, 1958, pp. 559-560) (Phillips [Birds of Arizona, 
1964, p. 159] believes some of Monniche’s specimens assigned to tolmiei 
are referable to O. philadelphia). W. W. Brown, Jr., found it at 
Boquete at 1200 m (Bangs, Proc. New England Zool. Club, vol. 3, 
1902, p. 61). The British Museum has a male and female collected by 
mance) labeled | Chiriqut’’ and “Volcan de Chirigqut.”, Whe California 
Academy of Sciences has 2 specimens identified as tolmiei collected at 
Barriles (1350 m, near Fl Volcan), Chiriqui on January 21 and Feb- 
ruary 4, 1931, and a male from the lowlands at Puerto Armuelles, 
taken November 7, 1929. My only encounter with this species was at 
El Volcan, where on February 12, 1955, J collected a male in low growth 
along a small quebrada running through a pasture still thick with felled 
tree trunks. 


294 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


All the records from the Canal Zone are quite old and were identified 
before mensural differences between this species and O. philadelphia 
were known. Stone (Proc. Acad. Nat. Sci. Philadelphia, vol. 70, 1918, 
p. 274) listed O. tolmiei from the Canal Zone. The Field Museum in 
Chicago has 2 females collected at Colon by J. F. Ferry on March 14 
and 16, 1908. The Smithsonian has a female collected by E. A. Gold- 
man at Gatun March 2, 1911, that fits the mensural characters of tolmiei 
(examined by W. Lanyon). 

Specimens collected by Monniche (Blake, op. cit.) on October 15 
and May 16 (to the extent correctly identified as tolmiei) would repre- 
sent the extreme dates for which this species is known in Panama. 

The specimens from Panama agree with those of the nominate race 
from California and Oregon in being brighter green above and brighter 
yellow below than monticola of Colorado and New Mexico. Cox (Auk, 
1973, pp. 190-191) describes occasional hybridization between this 
species and O. philadelphia in southwestern Alberta. In non-breeding 
plumage O. tolmie: cannot be safely separated in the field from O. 
philadelphia and must be identified by measurements, especially the 
relatively longer tail compared to wing; thus many identifications, even 
of specimens, made without determining wing and tail relationship (as 
described in Lanyon and Bull, Bird-Banding, 1967, pp. 187-194) are 
likely to be erroneous. 


GEOTHLYPIS TRICHAS (Linnaeus): Common Yellowthroat, 
Reinita Gargantiamarilla Comun 


Turdus Trichas Linnaeus, 1766, Syst. Nat., ed. 12, 1, p. 293. (Maryland.) 


Small, upper surface mostly buffy olive; throat (and on male much 
of undersurface) yellow; belly buff; male with black mask across fore- 
crown and face. 

Description.—Length 108-129 mm. Adult male, forecrown and side 
of head black, bordered by white except on sides of throat; crown buffy 
olive, becoming more greenish on back; wings dull green, with remiges 
edged yellowish green; tail yellowish green; throat and breast bright 
yellow fading to light buffy brown on belly; undertail coverts pale 
lemon yellow; bend of wing bright yellow; underwing coverts white. 

Adult female, forecrown light brown, fading to buffy olive on rest of 
upper surface; wings and tail as male; side of face dull tawny-olive; 
throat pale yellow, tinged with buff on breast and fading to light buffy 
brown on rest of undersurface; undertail coverts more yellowish. 

Migrant and winter visitor from the north. Rare in Bocas del Toro 


FAMILY PARULIDAE 295 


and Chiriqui and perhaps not found annually any farther east. The 
most easterly locations in Panama from which it has been reported is 
the EF] Real, Darién airstrip, where Ridgely saw 2 females on March 5, 
1981, and at Mandinga, San Blas, where | saw at least 2 in January and 
February of 1957. It usually forages near the ground, sometimes oc- 
curring with native yellowthroats. This species has been taken in 
Panama from October 15, when in 1967 a female was collected by the 
Gorgas Memorial Laboratory at Almirante, Bocas del Toro, to April 
24, when in 1963 R. Hinds collected a male at Almirante. Ridgely ob- 
served a male and a female at Tocumen marsh, eastern Province of 
Panama on April 29, 1976. Most Common Yellowthroats winter far- 
ther north, from Central America to the southern United States and in 
the West Indies. 

Geothly pis trichas has an extremely broad range across Mexico and 
North America and numerous races of varying degrees of distinction 
have been described. Dr. John W. Aldrich, who has studied this species 
extensively, very kindly examined the Panamanian specimens in the 
Smithsonian collection, and his identifications are followed here. 


GEOTHLYPIS TRICHAS BRACHIDACTYLUS (Swainson) 


Trichas brachidactylus Swainson, 1838 (1837?), Anim. Menag., p. 295. (Northern 
provinces of United States.) 


Characters —Large, extremely yellowish green above with pale 
flanks. 

Breeds in southern New England, eastern New York, and northern 
New Jersey. There are 3 specimens in the Smithsonian collection: 
USNM no. 459116 and no. 474689 from Juan Mina, Canal Zone, on 
January 9, 1955, and January 13, 1961, and USNM no. 469132 from 
Changuinola, Bocas del Toro, March 4, 1958. Another male specimen 
in the British Museum, no. 85.3.8.496, which agrees with the characters 
of this race, was collected by Arcé in 1870 at Volcan de Chiriqui, 
Chiriqui. 


GEOTHLYPIS TRICHAS PELAGITIS Braund and McCullagh 


Geothlypis trichas pelagitis Braund and McCullagh, 1940, Wilson Bull., 52, p. 118. 
(Eel Falls, 250 ft., Fox Bay, Anticosti Island.) 


Characters.—As large as brachidactyla, but darker greenish above 
and darker on flanks. 

Dr. Aldrich includes in this race the populations breeding from On- 
tario and northern Michigan through northern New England and mari- 


296 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


time Canada. A male, USNM no. 486554, taken at Almirante, Bocas 
del Toro, October 21, 1964, is of this race. 


GEOTHLYPIS TRICHAS MINNESOTICOLA Oberholser 
Geothlypis trichas minnesoticola Oberholser, 1948, Descr. New Races Geothlypis 
trichas (Linnaeus), p. 2. (St. Paul, Minnesota.) 

Characters ——Grayish green above, more rufescent on flanks and 
more orange-yellow on throat. 

Breeds in the northern Mississippi Valley from Iowa to Wisconsin 
and north to western Ontario and Manitoba. Specimens in the Smith- 
sonian are as follows: USNM no. 459115, a female from El Volcan, 
Chiriqui, March 14, 1955; USNM no. 462397, a male from Mandinga, 
San Blas, February 3, 1957; and USNM no. 469133, a male from 
Changuinola, Bocas del Toro, February 13, 1958. 


GEOTHLYPIS TRICHAS ROSCOE (Audubon) 

Sylvia roscoe Audubon, 1831, Orn. Bio. 1, p. 124. (not far from the River Mis- 

sissippi, Mississippi. ) 

Characters.—Small, paler, more yellowish green above; more rufes- 
cent on the flanks; more orange-yellow on throat. 

Breeds in the lower Mississippi Valley. E. A. Goldman collected a 
male, USNM no. 207165, at the Rio Indio, Canal Zone, on February 
13, 1911, that has been identified as this race. 


GEOTHLYPIS TRICHAS TYPHICOLA Burleigh 

Geothlypis trichas typhicola Burleigh, 1934, Proc. Biol. Soc. Washington, 47, p. 21. 

(Athens, Georgia.) 

Characters.—Brownish above and below, lighter colored, smaller. 

Breeds on the southeastern coastal plain from eastern Mississippi 
north to Virginia, exclusive of Florida. Dr. Aldrich has identified 3 
specimens from Almirante, Bocas del Toro, as belonging to this race: 
USNM no. 486553, a male collected April 24, 1963; AMNH no. 
785926, an immature male taken October 17, 1965; and a female from 
the Gorgas Memorial Laboratory collected October 15, 1967. 


GEOTHLYPIS AEQUINOCTIALIS CHIRIQUENSIS (Salvin): 
Masked Yellowthroat, Reinita Gargantiamarilla Chiricana 


Geothlypis chiriquensis Salvin, 1872, Ibis, p. 148. (Slopes of Volcan de Chiriqui, 
Panama. ) 
Small; upper surface bright yellowish green; undersurface yellow; 
male with black facial mask. 


FAMILY PARULIDAE 297 


Description.—Length 120-133 mm. Adult male, lores, forecrown, 
orbital ring, and side of face black; top of crown and line from rear of 
eye to nape, bordering black area, gray; hind crown and all of upper 
surface, including tail and wing feathers except for primaries, bright 
yellowish green; primaries dusky, with outer web edged bright yel- 
lowish green; sides and flanks greenish yellow; rest of undersurface, 
bend of wing, and underwing coverts bright yellow. 

Adult female, like male, but black replaced by gray, less extensive on 
side of face. 

Immature, like female, but with gray areas replaced by warbler 
green. 

Measurements.—Males (3 from Chiriqui), wing 58.1-61.5 (59.7), 
tail 46.7-51.2 (49.5), culmen from base 13.2-13.8 (13.6), tarsus 21.8- 
22.6 (22.1) mm. 

Females (2 from Chiriqui), wing 57.1-57.2 (57.2), tail 47.6-49.6, 
(48.6), culmen from base 13.7, tarsus 22.0-22.2 (22.1) mm. 

Resident. Found only in western Chiriqui in a few places around 
Volcan at 1050 to 1350 m, and in immediately adjacent southwestern 
Costa Rica, where it was first recorded by Skutch in 1964 ( Publ. Nutt. 
Orn. Club, no. 7, 1967, pp. 146-149). The Masked Yellowthroat in- 
habits lush meadows and marshy areas near streams; at La Lagunita, 
near El Volcan, I found a female perched in a spiny bush surrounded 
by heavy grass near a stream—habitat that Geothlypis trichas might 
choose. Although Griscom (Bull. Mus. Comp. Zool., vol. 78, 1935, p. 
369) called this species “very rare” it is actually very local but not un- 
common, usually found in loose colonies. Its voice is “a sweet twichee- 
teeweeo tweécheo, repeated four or five times, then a pause before 
starting again” (Ridgely, 1976, p. 301). In Costa Rica, Skutch (op. 
cit.) found them most vocal during May and June. Ridgely (i litt.) 
has heard them singing regularly in March in Chiriqui. 

On May 21, 1964, Skutch found a female working on a half-finished 
nest near Canas Gordas, Costa Rica. When completed, 4 days later, 
the nest was a bulky open cup composed of blades of lemon grass lined 
with long, smooth, brown fibers of unknown origin, situated about 45 
cm from the ground in a dense clump of lemon grass. The external 
dimensions were 10.1 by 14 cm in diameter by 8.9 cm high. The in- 
terior was 6.4 by 7.6 cm in diameter by 3.8 cm deep. The two eggs 
in this nest were laid May 26 and 27; they were “white, spotted with 
deep brown, pale brown, and pale lilac. The spots were largest and 
most crowded on the thick end of the egg and diminished toward the 
opposite pole, where they were few, small, and faint. The eggs mea- 


298 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


sured 20.0 by 13.9 and 19.6 by 14.1 mm. The shells had little gloss.” 
The eggs hatched after 15 days of incubation, and the nestlings were 
flesh colored, with a tinge of orange. Their natal down was a pale gray 
and by their third day the young had sprouted pinfeathers, but on that 
afternoon there was a heavy rainstorm that evidently killed the nest- 
lings. Skutch never found another nest. 

G. a. chiriquensis differs from other forms of G. aequinoctialts in its 
somewhat larger bill, duller dorsal coloration, and the greater extent of 
black on the forecrown. Despite the wide separation in range, the Chi- 
riqui bird is clearly a representative of aequinoctialis and Hellmayr’s 
(Field Mus. Nat. Hist., publ. 347, vol. 13, part 8, 1935, p. 440) listing 
of chiriquensis as a subspecies of aequinoctialis is probably justified, 
although most authors have kept them as separate species. It may also 
be noted that there is a similar, though less extreme, gap in the distri- 
bution of the races of Geothly pis semiflava, and geographical isolation 
is, therefore, not necessarily a compelling argument for specific dis- 
tinctness. 


GEOTHLYPIS SEMIFLAVA BAIRDI Ridgway: Olive-crowned 
Yellowthroat, Reinita Gargantiamarilla Coronioliva 


Geothlypis bairdi Ridgway (ex Nutting MS), 1884, in Nutting, Proc. U.S. Nat. 
Mus., 6 (1883), p. 398. (Los Sabalos, Nicaragua.) 


Small; upper surface yellowish olive-green; undersurface yellow; 
male has forecrown and facial area black. 

Description.—Length 111-125 mm. Adult male, forecrown, lores, 
and side of face to lower edge of throat black; rest of upper surface, 
including tail and all of wings except primaries, yellowish olive-green; 
primaries dusky with outer web edged olive-green; sides and flanks 
olive-green; rest of undersurface, bend of wing, and underwing co- 
verts bright lemon yellow. 

Adult female, like male, but black replaced by olive- green, more yel- 
lowish on forehead and superciliary; olive wash on undersurface more 
extensive. 

Immature, like female, but buffier below. 

Measurements.—Males (10 from Bocas del Toro and Costa Rica), 
wing 57.2-62.0 (59.2), tail 42.7-52.4 (47.6), culmen from base 13.8- 
16.1 (14.9), tarsus 20.1-22.9 (21.7) mm. 

Females (7 from Bocas del Toro, Costa Rica, and Nicaragua), wing 
54.2-57.0 (55.7), tail 42.6-47.3 (45.1), culmen from base 13.9-15.2 
(14.7), tarsus 20.1-21.8 (21.2) mm. 


FAMILY PARULIDAE 299 


Resident. Common in the lowlands of western Bocas del Toro, and 
north on the Caribbean slope to southern Honduras. In Panama this 
species inhabits rank grass, bamboo thickets, shrubbery, and the bor- 
ders of reedy swamps. I have collected it at Almirante and Changui- 
nola; H. von Wedel collected it also at Isla Grande, in the Sixaola River 
near Victoria, about 32 km northwest of Almirante (Peters, Bull. Mus. 
Comp. Zool., vol. 71, 1931, p. 337). G. semiflava was not recorded from 
Panama prior to 1926, when on February 14 Kennard collected a male 
and female at Almirante (Kennard and Peters, Proc. Boston Soc. Nat. 
- Hist., vol. 38, 1928, p. 460). Its behavior is like other yellowthroats, 
and I have found it hard to distinguish from migrant G. trichas until I 
have had the specimen in hand. The song is a loud, sweet warble, some- 
what suggestive of the Gray-crowned Yellowthroat and Orchard 
Oriole (Ridgely, 1976, p. 301). Three nests collected in early April 
of 1970 and 1971 in Costa Rica, now at the Western Foundation of 
Vertebrate Zoology, were each found in pasture. They were open cups 
made of wide dead strips of grass and lined with finer grass. Lloyd 
Kiff (pers. comm.) describes the eggs as ‘“‘white with sparse springling 
of fine black and dark brown dots, mostly at the large ends. Similar to 
G. trichas eggs, but lacking the scrawls that are present on many ex- 
amples of the latter.” Measurements of 5 eggs average 19.47 14.05 
mm. Two eggs form a clutch. 

Eisenmann points out that the yellowish (rather than whitish or 
brownish) abdominal area separates it from G. trichas in all plumages, 
and males can be distinguished from all members of the genus known 
from Panama by the more extensive black on the crown and lack of 
gray or white on the head. 


GEOTHLYPIS POLIOCEPHALA RIDGWAYI (Griscom): Gray- 
crowned Yellowthroat, Reinita Gargantiamarilla Carbonera 


Chamaethlypis poliocephala ridgwayi Griscom, 1930, Proc. New England Zool. 
Club, 12, p. 7. (Boruca, southwestern Costa Rica.) 


Small; crown gray; rest of upper surface grayish olive-green; un- 
dersurface yellow. 

Description.—Length 115-131 mm. Adult male, lores and area just 
below eye black; crown gray; side of face and rest of upper surface, 
grayish olive-green, brighter, more yellowish on rump, upper tail co- 
verts, and wings; undersurface, bend of wing, and underwing coverts 
yellow, fading into buffy whitish on abdomen and to light buffy olive 
on sides and flanks. 


300 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


Adult female, like male, but gray on crown less extensive, rest of up- 
per surface slightly browner than male; black area less intense, and 
yellow undersurface paler. 

Measurements.—Males (8 from Chiriqui and Costa Rica), wing 
53.5-57.0 (54.8), tail 51.7-61.1 (56.3), culmen from base 11.8-13.2 
(12:5)); tarsus 20.4-23:20(2107\) arama, 

Females (5 from Chiriqui and Costa Rica), wing 50.0-57.7 (53.3), 
tail 54.9-62.1 (57.7), culmen from base 11.7-13.2 (12.5), tarsus 21.0- 
2Z,81(20S i coma: 

Resident. Uncommon and local in the foothills and highlands of 
western Chiriqui from 690 to 2250 m (Ridgely, 1976, p. 301), and 
west to the Térraba Valley in southwestern Costa Rica. Other races 
are found from extreme southern Texas through Middle America. 
This yellowthroat inhabits overgrown and abandoned pastures and 
brushy fields. Monniche collected specimens at Lérida and Quiel on 
the Volcan de Chiriqui (Blake, Fieldiana: ‘Zool., vol 36;me}aeoae 
p. 560) and W. W. Brown, Jr., collected it also at Boquete (Bangs, 
Proc. New England Zool. Club, vol. 3, 1902, p. 61). The only place I 
found this species was at El Salto (1200 m), near Boquete, where on 
March 17 and 18, 1960, I collected a male and female in a field of 
bracken and grass. The song, which is sometimes delivered from an 
exposed perch as high as a tree branch or a wire, is “somewhat similar 
to semiflava and has a number of undistinctive vireo-like phrases strung 
together. It 1s not so long as that of semiflava, does not ramble up and 
down so markedly, and lacks a slur at the end although one may be 
thrown in elsewhere” (Slud, Bull. Amer. Mus. Nat. Hist., vol. 128, 
1964, p. 330). This species also has at least a half dozen calls. Rowley 
(Condor, 1962, p. 262) found the nest of the nominate race near 
Cuernavaca, Mexico. It was in dry grass bordering a cleared field and 
was made entirely of dried grasses; it contained 3 eggs. 

Eisenmann (Auk, 1962, pp. 265-267) reviews all characters of this 
species, formerly placed in the monotypic genus Chaemethlypis, and 
concludes that it is not allied to cteria but is properly placed in the 
genus Geothly pis, the only distinction between it and other members of 
the genus being the larger bill. It is “simply a yellowthroat of the drier, 
more sterile uplands, as distinct from the marshes or borders of wet 
places preferred by other yellowthroats. The drier niche, with the con- 
sequent difference in diet, doubtless accounts for the heavier bill, which 
is presumably useful in feeding on harder-bodied or larger insects (and 
possibly on seeds) than are normally taken by yellowthroats of water 


FAMILY PARULIDAE 301 


edges, which have more soft-bodied larval forms available. The re- 
duction of the black mask also seems correlated with a drier environ- 
ment.” 


ICTERIA VIRENS VIRENS (Linnaeus): Yellow-breasted Chat, 
Reinita Arriera 


Turdus virens Linnaeus, 1758, Syst. Nat., ed. 10, 1, p. 171. (South Carolina.) - 


Medium size; upper surface yellowish olive; abdomen and undertail 
coverts white; rest of undersurface yellow. 

Description.—Length 151-174 mm. Adult (sexes alike), forehead 
gray; superciliary and lower rim of eye white; lores black; upper sur- 
face and wing coverts yellowish olive; remiges and rectrices dusky with 
outer webs edged light yellowish olive; auricular region gray; thin 
white stripe on side of throat; throat to belly bright yellow; flanks gray; 
rest of undersurface white; bend of wing and underwing coverts yellow. 

Measurements.—Males (10 from the breeding range, taken in May), 
wing 71.0-78.0 (75.5), tail 69.6-77.5 (74.4), culmen from base 14.2- 
17.0 (15.3), tarsus 24.3-27.0 (25.7) mm. 

Females (10 from the breeding range, taken in June), wing 71.8- 
80.0 (76.5), tail 66.5-83.4 (76.2), culmen from base 14.9-16.1 (15.4), 
tarsus 24.7-27.8 (26.0) mm. 

Winter visitor from the north. Uncommon in the lowlands of west- 
ern Bocas del Toro, where it has been recorded at Almirante and at the 
mouth of the Rio Teribe on the Rio Changuinola. The first specimen 
known from Panama was collected at Almirante by H. von Wedel on 
January 16, 1929, but banding results suggest that it is not rare there: 
in 1963, 16 were netted October 1-31 and 3 were netted April 17-21 
WEettinvand Olson) Carib. Journ: Sci.) vol.\3; no. 4, 1963, p, 195,). 
Banding also indicates that Yellow-breasted Chats return to the same 
site in succeeding years. One banded at Almirante April 17, 1963, was 
recaptured there October 17 and 18, 1963, and again October 22 and 
Movember 11965, (Loftim et al.. Bird-Bandimeg, 1967, \p. 152). The 
only report of this species elsewhere in Panama, and the most southern, 
is a sighting of 1 on Cerro Campana (far to the east of any other record 
and the only one from the Pacific slope) on September 14, 1961, by 
Richard Ryan and Ned Boyajian (Ridgely, 1976, p. 301). In Panama, 
as in its breeding range, the chat usually stays well hidden in dense 
undergrowth of brushy areas and woodland borders. 


302 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


WILSONIA CITRINA (Boddaert): Hooded Warbler, 
Reinita Encapuchada 


Muscicapa citrina Boddaert, 1783, Tabl. Planches Enlum., p. 41. (Louisiana. ) 


Small; face yellow; hindcrown, sides of neck, and throat black; rest 
of upper surface olive-green; rest of lower surface yellow. 

Description.—Length 118-132 mm. Adult male, forehead, fore- 
crown, and side of face to auriculars bright yellow; loral spot, re- 
mainder of crown, nape, sides of neck, throat, and upper breast black; 
rest of upper surface olive-green; remiges dusky, with outer web 
edged olive-green; two outer pairs of rectrices nearly all white on 
inner web, dusky on outer web; next pair with white spot on inner 
web; remaining rectrices dusky, with outer web edged olive-green; 
rest of undersurface yellow, fading to very pale yellow on under- 
tail coverts; bend of wing yellow; underwing coverts white. 

Adult female, upper surface olive-green; wings and tail as male, but 
duller; lores, superciliary, side of face bright yellow, with auricular 
region tinged greenish yellow; undersurface bright yellow, becoming 
paler on undertail coverts. 

Measurements.—Males (10 from the breeding range, taken in May 
and June), wing 62.1-70.0 (66.6), tail 53.2-58.1 (55.3), culmen from 
base 10.7-12.8 (11.6), tarsus 17.9-20.5 (19.3) mm. 

Females (10 from the breeding range, taken in April, May, and 
June), wing 60.5-65.0 (62.5), tail 50.8-56.5 (54.5), culmen from base 
10.5-12.0 (11.2), tarsus 18.6-19.8 (19.1) mm. 

Winter visitor from the north. Breeds in eastern United States; 
winters through Middle America to central Panama. Rare in the low- 
lands of western Bocas del Toro (Almirante) and the Canal Zone area. 
An adult male was reported seen by N. G. Smith in “late September” 
1964 and another in Panama City in his garden on May 2, 1968 (in litt. 
to Eisenmann). McLeannan evidently encountered this species at least 
twice in what became the Canal Zone, since specimens collected by him 
are now in the British Museum (Salvin and Godman, Biol. Centr.- 
Amer., Aves, vol. 1 (pt. 12), 1881, p. 167) and the American Museum 
of Natural History (Lawrence, Ann. Lyc. Nat. Hist. New York, vol. 
8, 1863, p. 6). More recently in the Canal Zone, Gale and West (im 
litt.) banded, photographed, and released a male at Corozal on October 
27,1973, and T. A. Imhof (in litt.) found 1 at Caflo Saddle on Decem- 
ber 29, 1942. A male was seen on Barro Colorado Island on September 
24, 1951, and another on January 5, 1977 (Willis and Eisenmann, 
Smiths. Contrib. Zool. 291, 1979, p. 27). In Bocas del Toro only 3 or 
4 were banded in two seasons (October 12-30) at Almirante (Loftin, 


FAMILY PARULIDAE 303 


in litt. to Eisenmann). Extreme dates from Panama (fide Eisenmann) 
are September 24-May 2. Willis (Living Bird, 1966, p. 209) considers 
the Hooded Warbler to be an ecological competitor of the Spotted Ant- 
bird (Hylophylax naevioides) and suggests that this is why the warbler 
is locally rare or absent as a winter resident wherever the antbird occurs. 

A male collected at Almirante by D. L. Hicks on October 25, 1964, 
weighed 8 g. 


WILSONIA PUSILLA (Wilson): Wilson’s Warbler, Reinita de Wilson 


Small; mid-crown black; upper surface bright olive-green; forehead 
and undersurface yellow. 

Description.—Length 103-115 mm. Adult male, forehead, super- 
ciliary, and orbital ring bright yellow; center and rear of crown shin- 
ing black; auriculars and rest of upper surface bright olive-green; 
wings and tail dusky, with wing coverts tipped and rectrices and remiges 
edged olive-green; undersurface, bend of wing, and underwing coverts 
bright yellow. 

Adult female, like male, but black on crown less extensive or absent. 

Wilson’s Warbler is an abundant winter resident in the highlands of 
Chiriqui, where, at Fl Volcan, I saw as many as a dozen together, on 
March 13, 1954. In the highlands and foothills farther east it becomes 
progressively less common. It is seen regularly in small numbers above 
Santa Fé, Veraguas, but in central Panama there are only 3 sight rec- 
ords, all of males: 1 on Cerro Campana on September 14, 1961 (R. 
Ryan), and single individuals on Cerro Azul on December 30, 1967, 
and December 28, 1973 (Ridgely, 1976, p. 302). Panama is the south- 
ern limit of this species’ range in winter. Panama records (fide Eisen- 
mann) run from September 10 to May 16 (both specimen records). 

Leck ( Bird-Banding, 1975, p. 202) found that 15 Wilson’s Warblers 
mist netted at Cerro Punta, Chiriqui, during September and October 
1967 had an average weight of 7.6 g and 12 netted there in March 1968 
averaged 8.0 g. 

I have seen Wilson’s Warblers feeding in herbaceous growth near 
the ground and as high as 10 m up in second growth. They are often 
common in coffee plantations. This is the most numerous and con- 
spicuous wintering wood warbler in the Chiriqui highlands. Ridgely 
(in litt.) has seen up to 30-35 in a day in Chiriqui and in the Costa Rica 
highlands as well. 

Itisenmann (im litt.) noted that in the Chiriqui highlands Wilson’s 
Warblers favor semi-open or lightly wooded areas. He found them 
usually moving about in the mixed bands of wandering warblers and 


304 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


tanagers. Ordinarily, only 1 Wilson’s Warbler was in each band, but 
occasionally there were 2 or 3. They were noisy, constantly giving a 


chip-chip-chip-chip or tsip-tsip-tsip. 


WILSONIA PUSILLA PILEOLATA (Pallas) 


Motacilla pileolata Pallas, 1811, Zoogr. Rosso-Asiat., 1, p. 497. (Kodiak Island, 
Alaska.) 


Characters.—Forehead light cadmium; upper surface paler green. 

Measurements.—Males (10 from Chiriqui), wing 55.0-61.0 (57.1), 
tail 47.8-52.0 (50.0), culmen from base 9.7-11.3 (10.7), tarsus 17.2- 
BOLO USS) i montde 

Females (10 from Chiriqui), wing 55.0-59.4 (56.0), tail 45.8-48.6 
(47.5), culmen from base 9.2-11.4 (10.2), tarsus 16.8-18.8 (18.0) mm. 

Winter visitor from the north, breeding in the Rockies and most of 
western Canada and northwestern United States. Abundant in the 
highlands of western Chiriqui, less numerous farther east. Monniche 
(Blake, Fieldiana: Zool., vol. 36, no. 5, 1958, p. 560) collected this race 
on the Volcan de Chiriqui between 1560 and 2700 m. W. W. Brown, 
Jr., found it at Boquete as low as 1200 m (Bangs, Proc. New England 
Zool. Club, vol. 3, 1902, p. 61); I have seen the species there from 1260 
to 2610 m. Monniche collected specimens of this race from September 
17 to May 16. Griscom (Bull. Mus. Comp. Zool. 78, 1935, p. 369) 
considered this race fairly common in Veraguas and listed it from the 
Canal Zone. 


WILSONIA PUSILLA CHRYSEOLA Ridgway 


Wilsonia pusilla chryseola Ridgway, 1902, Bull. U.S. Nat. Mus., 50, pt. 2, pp. 705, 
714, (Red Bluff, California.) 


Characters.—Upper surface suffused with yellow, lighter than any 
other race. 

Measurements.—Males (10 from North America, taken in April and 
May), wing 52.0-58.5 (55.0), tail 44.1-49.9 (48.2), culmen from base 
96-1 1.9) (10:5))) tarsus l5:9219.9 (1722) amae 

Females (10 from North America, taken in May), wing 50.8-55.5 
(52.8), tail 44.5-49.9 (47.1), culmen from base 9.7-11.3 (10.8), tarsus 
17.3-19.0 (18.2) mm. 

Winter visitor from the north, breeding in the Pacific coast region of 
the United States and Canada. Rare in the highlands of western Chiri- 
qui. I know of only 4 specimens of this well-marked race that have 
been taken in Panama. Arcé collected a male on the southern slope of 


FAMILY PARULIDAE 305 


the Volcan de Chiriqui in 1870. The specimen is now in the British 
Museum and is presumably the one to which Griscom was referring 
when he listed “fone record” for Panama (Bull. Mus. Comp. Zool., 78, 
1935, p. 369). C. Lynn Hayward collected 3, a male and 2 females, on 
Cerro Punta between March 6 and 14, 1962; they are now in the 
Monte L. Bean Life Sciences Museum at Brigham Young University, 
Provo, Utah. 


WILSONIA PUSILLA PUSILLA (Wilson) 


Muscicapa pusilla Wilson, 1811, Amer. Ornith., 3, p. 103, pl. 26, fig. 4. (Southern 
New Jersey.) 


Characters.__Forehead lemon chrome; upper surface darker green. 

Measurements.—Males (10 from North America, taken in May), 
wing 53.3-56.0 (54.5), tail 44.3-49.3 (47.1), culmen from base 9.6- 
11.4 (10.3), tarsus 17.2-18.4 (17.8) mm. 

Females (10 from North America, taken in May), wing 50.8-55.1 
(54.0), tail 44.0-49.4 (46.7), culmen from base 10.1-11.4 (10.8), tarsus 
16.1-19.5 (18.1) mm. 

Winter visitor from the north, breeding in northwestern United 
States and eastern Canada. Fairly common in the highlands of western 
Chiriqui. I have collected individuals of this race at El Volcan from 
1320 to 1560 m, at Santa Clara (1080 m), and on Cerro Punta (2010 


m). 


WILSONIA CANADENSIS (Linnaeus): Canada Warbler, 
Reinita de Canada 


Muscicapa canadensis Linnaeus, 1766, Syst. Nat., ed. 12, 1, p. 327. (Canada.) 


Small; upper surface gray; undersurface yellow, with necklace of 
black streaks on breast of male. 

DescriptionLength 113-127 mm. Adult male, forehead to top of 
crown black, with feathers tipped gray; rest of upper surface, includ- 
ing wings and tail, gray; superciliary to eye and orbital ring yellow; 
lores, auriculars, side of throat, and spots on breast black; undersur- 
face yellow, to undertail coverts, which are white; underwing coverts 
white. In winter upper surface tinged with olive. 

Adult female, like male, but black areas reduced, paler, or absent; 
yellow on undersurface slightly paler. 

Measurements——Males (10 from the breeding range, taken in May 
and June), wing 64.0-69.5 (65.5), tail 51.2-54.6 (53.0), culmen from 
base 10.5-12.5 (11.5), tarsus 17.4-20.0 (18.6) mm. 


306 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Females (10 from the breeding range, taken in May and June), wing 
59.2-63.3 (61.5), tail 49.3-52.5 (51.4), culmen from base 10.4-12.0 
(11.6), tarsus 17.2-18.8 (18.1) mm. 

Migrant and winter visitor from the north. Fairly common through- 
out on migration, in small numbers up to the highlands of western 
Chiriqui as high as 1620 m (Blake, Fieldiana: Zool., vol. 36, no. 5, 1958, 
p. 560), and on the Pearl Islands (Rendahl, Ark. Zool, vol. 13, no. 4, 
1920, p. 48). The Canada Warbler has been recorded in Panama from 
September 3 to May 19 (Eisenmann, Smiths. Misc. Coll., vol. 117, no. 
5, 1952, p. 52; Willis and Eisenmann, Smiths. Contrib. Zool. 291, 1979, 
p. 27); it is most numerous from late September to early October and 
in late April and early May. On May 1, 1980, Ridgely counted at least 
18 (75% of them males) at Galeta Island, Canal Zone. In 1963, 94 
were banded at Almirante, Bocas del Toro, between September 18 and 
October 28 (Loftin, im litt.). Willis (Living Bird, 1966, p. 209) saw 
12 on Barro Colorado Island on October 3, 1960; he found that the 
Canada Warbler regularly followed swarms of army ants there with as 
many as 5 individuals in attendance at one swarm. In Panama it 
usually occurs in the understory of forest and woodland, often at edge, 
but only infrequently out in clearings, and then only when they are 
heavily overgrown. 

This species is recorded in winter from British Honduras south to 
central Peru and northern Brazil, but is scarce at that season north of 
South America, its primary wintering area. In winter it has been re- 
corded from Cerro Campana, Province of Panama, and from the Canal 
Zone (Ridgely, op. cit). One banded at Curundu in the Canal Zone on 
February 13, 1964, and retaken there 6 weeks later on March 29 
(Loftin et al., Bird-Banding, 1966, pp. 42-43) was probably a winter 
resident. I also have a sight record for Chilar, western Colon, on Feb- 
ruary 19, 1952. Two sightings that probably represent wintering indi- 
viduals in Chiriqui were an immature female at Fortuna on March 3, 
1976, and a male near Volcan (Dos Rios Hotel) on March 20, 1979, 
both observed by Ridgely. 


SETOPHAGA RUTICILLA (Linnaeus): American Redstart, 
Candelita Americana 


Motacilla ruticilla Linnaeus, 1758, Syst. Nat., ed. 10, 1, p. 186. ( Virginia.) 
Small; male black with orange on remiges, rectrices, and sides; fe- 


male with gray crown, rest of upper surface green, undersurface white 
with yellow on sides. 


FAMILY PARULIDAE 307 


Description.—Length 113-126 mm. Adult male, entire upper sur- 
face, throat, and breast shining black; wings black with basal half of 
remiges pinkish orange; central pair of rectrices black, all others pink- 
ish orange on basal half, black on distal half; sides and flanks orange; 
rest of undersurface white; underwing coverts pinkish orange. 

Adult female, crown gray; rest of upper surface to upper tail co- 
verts buffy olive; upper tail coverts blackish; remiges blackish brown 
with yellow, base and outer webs edged buffy olive; central pair of 
rectrices blackish brown, all others with basal half wholly or partially 
light yellow and remainder blackish brown; orbital ring white; side of 
head gray; sides and flanks yellow; rest of undersurface white; under- 
wing coverts pale yellow. 

Immature male, like female, but sides of breast tinged pinkish orange. 

Measurements.—Males (10 from the breeding range, taken in May) 
wing 61.8-66.0 (64.1), tail 52.3-57.4 (54.7), culmen from base 10.0- 
11.6 (10.8), tarsus 15.0-17.2 (16.2) mm. 

Females (10 from the breeding range, taken in May and June), 
wing 58.0-63.8 (60.4), tail 51.4-56.2 (53.4), culmen from base 10.5- 
120 (11), tarsus 15.2-18.3 (16:3) mm. 

Migrant and winter visitor from the north. Fairly common through- 
out on migration, less common in winter. The American Redstart has 
been recorded as high as 1650 m on the Volcan de Chiriqui (Blake, 
Fieldiana: Zool., vol. 36, no. 5, 1968, p. 561); I have also seen it in 
coastal mangroves, at Tonosi, Los Santos, and Almirante, Bocas del 
Toro, for example. Aldrich (Scient. Publ. Cleveland Mus. Nat. Hist., 
vol. 7, 1937, p. 125) saw it on several occasions in forests on the west 
side of the Azuero Peninsula, Veraguas, where it was part of mixed 
flocks of migrant North American warblers. I have found redstarts in 
high forest on Isla Coiba, and W. W. Brown, Jr. (Thayer and Bangs, 
Bull. Mus. Comp. Zool., vol. 46, 1905, p. 156) collected a female on San 
Miguel in the Pearl Islands on March 2, 1904. The British Museum 
has specimens collected by H. J. Kelsall at sea 160 to 190 km south of 
Panama, taken between August 28 and October 2, 1924. 

The American Redstart has been found in Panama as early as Au- 
eosel/ (kisenmann, Smiths. Misc. Coll, volull7, no. 5; 1952) sp) 52) 
and as late as May 11 (Ridgely, 1976, p. 302); most depart by late 
April. In Eisenmann’s experience their numbers vary greatly from 
year to year; in some seasons few are noted even in fall migration. One 
banded near Curundu, Canal Zone, on December 8, 1963, was re- 
captured there September 27, 1965 (Loftin et al., Bird-Banding, 1967, 
p. 152). In Panama this species is found in woodlands and borders. It 


208 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


is less numerous in the highlands than at lower elevations. Ridgely (in 
litt.) has found it more common in mangroves than any other habitat 
in winter. Eisenmann has seen it on migration in the semi-arid, rather 
scrubby western part of the Province of Panama near the coast at 
Playa Coronado, on August 19-26, 1947, (2-4 birds daily) and at Santa 
Clara Beach. 


MYIOBORUS MINIATUS (Swainson): Slate-throated Redstart, 
Candelita Gargantigris 


Setophaga mimata Swainson, 1827, Philos. Mag., new ser., 1, fasc. 5, p. 368. 
(Morelia, Michoacan. ) 


Small; upper surface and throat dark gray, with chestnut patch in 
center of crown; undersurface yellow. 

Description.—Length 118-128 mm. Adult (sexes alike), upper sur- 
face, wings, head and throat slate color (throat dark, sometimes black) ; 
center of crown chestnut; tail black, with outer pair of rectrices nearly 
all white, second pair with distal half white, third pair white only at 
tip; undertail coverts white; rest of undersurface yellow; underwing 
coverts white. 

Juvenile, head, back, rump, and foreneck sooty slate; upper breast 
and sides washed with dull rufous; lower breast, abdomen, and under- 
tail coverts cinnamon; wings and tail like adults. 

The Slate-throated Redstart is very common in woodlands, forests, 
and even borders and gardens adjacent to clearings in the highlands of 
Chiriqui and Veraguas and in eastern Darién. It moves about like 
Seto phaga ruticilla, opening and shutting its tail both while perched and 
in flight, so that the white on the outer feathers is conspicuous. I have 
found these birds high in the tree crowns, in the undergrowth, and on 
the ground, where they occasionally drop when pursuing prey. Leck 
(Living Bird, 1971, p. 91) found that at Cerro Punta, Chinqnw aiey 
spent 45 percent of their time in the upper middle section of trees and 
another 40 percent in the lower middle. Their food is almost exclu- 
sively insect material, but Skutch (Pac. Coast Avif. no. 31, 1954, p. 
358) has observed them eating the little white protein bodies that are 
found at the base of the petioles of leaves on Cecropia trees. The Slate- 
throated Redstart sings throughout the year, although less often 
between October and January; it has “several usually squeaky or 
sibilant songs, sometimes suggestive of American Redstart, tseeweech, 
sweeswee, sweech-sweechee, or tseeoo-tseeoo, cheewee-cheewee-tsee”’ 
(Ridgely, 1976, p. 303). 

This species is found from northern Mexico through Middle Amer- 


FAMILY PARULIDAE 309 


ica to Guyana, northern Brazil, and Bolivia. In Panama the birds of 
the western highlands belong to the race aurantiacus and those of 
Darién are of the race ballux. 


MYIOBORUS MINIATUS AURANTIACUS (Baird) 
Setophaga aurantiaca Baird, 1865, Rev. Amer. Birds, 1, p. 261. (Dota Mountains, 
San José and Barranca, Costa Rica.) 


Myioborus miniatus acceptus Bangs, 1908, Proc. New England Zool. Club, 4, p. 
30. (Boquete, 4000 ft., Volcan de Chiriqui, Panama.) 


Characters.—Chestnut crown patch bordered by black stripes; un- 
dersurface slightly darker yellow. 

Measurements.—Males (10 from Chiriqui), wing 61.0-68.8 (63.2), 
tail 57.1-63.7 (61.3), culmen from base 10.6-13.0 (11.9), tarsus 16.3- 
19.6 (17.8) mm. 

Females (10 from Chiriqui), wing 58.0-67.5 (60.9), tail 54.2-62.8 
(S62) culmen from base 10:6-12.2 (11.3), tarsus 16.7-18.6 (17.7) 
mm. 

Resident. Very common in the highlands of Chiriqui and Veraguas. 
Found also in the Cordillera de Talamanca of eastern Costa Rica; 
Monniche collected it on the Volcan de Chiriqui between 1590 and 
1890 m (Blake, Fieldiana: Zool., vol. 36, no. 5, 1958, p. 561) and W. W. 
Brown, Jr., found it from 1200 to 1350 m at Boquete, Chiriqui, during 
January and February of 1901 (Bangs. Proc. New England Zool. 
Club, vol. 3, 1902, p. 61). It is common in the regenerating woodland 
around the Volcan lakes (1200 m). Griscom (Bull. Mus. Comp. Zool., 
vol, 78, 1935, p. 369) said it occurred above 1050 m. At Cerro Punta, 
Chiriqui, I found it rather uncommon at 2100 m during March 1955; I 
believe this must be about the upper limit of its range. The only locality 
in Veraguas from which I know of a specimen is Calovévora, where 
Arcé collected a male, now in the British Museum, on 1867. In early 
January 1974, Ridgely (1 litt.) found it fairly common along the road 
north of Santa Fé (900-1160 m). 

On March 3, 1954, I saw that males were in pursuit of females at 
EI Volcan, Chiriqui, and in 1960 I found that they were always in pairs 
there on February 24. In 1965 I collected a male in breeding condition 
at El Volcan on February 27. Further details of breeding behavior 
come from Costa Rica, where Skutch (Pac. Coast Avif. no. 31, 1954, 
pp. 357-365) has found several nests, always on the ground. Most that 
Skutch found were in a cranny in a bank, but others were placed on a 
fallen log amid dense vegetation and on a steep slope, also well sheltered 
by vegetation. The nest is a roofed structure composed of straws, dried 


310 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


grass, and fibrous rootlets, and requires 3 to 5 days of construction. 
Skutch found nests between March 30 and mid-May. 

A week usually elapsed between completion of the nest and laying 
of the first egg. Eleven nests that Skutch found contained 3 eggs or 
nestlings, 2 contained 2 eggs each. “The eggs were white or dull white, 
speckled and blotched with some shade of brown, ranging from bright 
brown to chocolate. The pigment was heaviest in a wreath about the 
point of greatest transverse diameter, or in a cap covering the large 
end... The measurements of 20 eggs averaged 17.5 by 13.4 milli- 
meters” (Skutch, of. cit., p. 362). The incubation period varied from 
13 to 15 days, and at hatching the nestlings were pink-skinned and 
blind. By 9 or 10 days of age they were well feathered, and left the nest 
at 12 to 14 days. By mid-July the young nearly resemble their parents 
and in October they appear to be mated. While observing the nests 
of this species, Skutch found that parents would sometimes feign in- 
jury, fluttering on an open piece of ground near their nest. 

On April 23, 1961, Eisenmann at Nueva Suiza, Chiriqui, ca. 1800 m, 
saw a pair carrying food to a domed nest on the ground. During this 
month he heard birds singing tsireé-tsireé-tsireé, tsee-tsee-tsee (lasting 
about 2 seconds); in late February he noted a different, rather fast 
tseéoo-tseéoo, cheéwe p-cheéwee-tsee, somewhat suggestive of a song 
of the American Redstart, but weaker and rather less sweet. In Sep- 
tember Eisenmann heard the song quoted by Ridgely. It also gives a 
call, tit-tit-tit-tit, as it flicks open the tail, snappier and louder than a 
similar call of the Collared Redstart. 


MYIOBORUS MINIATUS BALLUX Wetmore and Phelps 


Myioborus miniatus ballux Wetmore and Phelps, 1944, Proc. Biol. Soc. Wash- 
ington, 57, p. 11. (1,600 m., near Queniquea, Tachira, Venezuela.) 


Characters.—White on outer rectrices more extensive than in M. m. 
aurantiacus; chestnut crown patch not bordered with black; lighter 
yellow below. 

A male collected February 22, 1964, at Cerro Mali, Darién, had the 
iris slightly reddish brown (very dark); bill black; tarsus, toes, and 
claws fuscous-black. 

Measurements.—Males (10 from Darién and Colombia), wing 55.0- 
66.1 (62.0), tail 51.3-62.2 (56.8), culmen from base 10.4-12.9 (11.5), 
tarsus 14.9-18.4 (16.9) mm. | 

Females (10 from Darién and Colombia), wing 57.0-64.0 (60.5), 


FAMILY PARULIDAE QLe 


tail 49.9-59.1 (56.0), culmen from base 10.6-11.8 (11.4), tarsus 16.3- 
18.2 (17.4) mm. 

Resident. Very common in forest in eastern Darién between 1050 
and 2100 m. It has been collected at Cerro Mali, Cerro Cana, Cerro 
Tacarcuna, Cerro Pirre, and Alturas de Nique. A male that I collected 
on Cerro Mali on February 26, 1964, was in breeding condition and 
had somewhat worn plumage. E. A. Goldman’s notes from Cerro Pirre 
in 1912 mention a female collected April 18 “with egg ready to lay.” 
On April 27 he says “‘this species is nesting.” 

An individual seen in cloud forest on Cerro Jefe, eastern Province of 
Panama in July 1973 by D. Hill (Ridgely, 1976, p. 303) may have 
been of this race, which ranges south through the Andes of Colombia 
and western Venezuela at least as far as northwestern Ecuador. 

AY H. Miller (Univ. Calif. Publ. Zool., vol no. 1, 66, 1963, p. 45) 
has described habitat, behavior, nesting, and weights (8 males, 9-10 g; 
female, 9.3 g) of M. m. ballux in the western Andes of Colombia at 
San Antonio. 


MYIOBORUS TORQUATUS (Baird): Collared Redstart, 
Candelita Collareja 


FIGURE 24 


Setophaga torquata Baird, 1865, Rev. Amer. Birds, 1, p. 261. (San José, Costa 
Rica.) 


Small; upper surface dark gray with reddish brown crown; sides of 
face and undersurface yellow, with black band across upper breast. 

Description.—Length 121-133 mm. Adult (sexes alike), forecrown, 
sides of crown, and nape black; center of crown rufous; upper surface 
slate color; wings and tail blackish, with white extensive on outer two 
pairs of rectrices; forehead, lores, superciliary, side of head, and under- 
surface yellow, with black band across chest; underwing coverts a mix- 
ture of light yellow and gray. 

Immature, upper surface gray; forehead, lores, lower cheeks, and 
throat pale yellow flecked with gray; broad gray band across breast; 
lower breast and belly yellow; brightest on flanks, fading to whitish on 
center of abdomen. 

A male taken March 5, 1965, at Volcan de Chiriqui, Chiriqui, had the 
iris mouse brown; bill black; tarsus and toes fuscous-brown; claws 
black. 

Measurements.—Males (10 from Chiriqui), wing 63.9-70.1 (66.7), 


312 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


tail 56.8-63.8 (60.1), culmen from base 10.3-12.4 (11.4), tarsus 17.9- 
20.3 (19.4, average of 9) mm. 

Females (10 from Chiriqui), wing 61.1-64.5 (62.0), tail 54.7-59.3 
(57.8), culmen from base 9.6-11.8 (10.4, average of 9), tarsus 18.4- 
199" (C1910) samt 


Ficure 24.—Collared Redstart, Candelita Collareja, Myioborus torquatus. 


Resident. Common in highlands of Chiriqui, adjacent Bocas del 
Toro, and in Veraguas, in forests and woodland edges. It is also found 
in the mountains of Costa Rica. Monniche found it up to 2700 m on the 
Volcan de Chiriqui, higher than M. miniatus occurs (Blake, Fieldiana: 
Zool., vol. 36, no. 5, 1958, p. 561). In Chiriqui it is usually found above 
1800 m, sometimes down to 1050 (at the Fortuna Dam site); in Vera- 
guas (Chitra, Santa Fé, Cerro Tute) it regularly comes down to 1050- 
1200 m (Ridgely, 1976, p. 303). M. torquatus has been collected com- 
monly in Chiriqui on the Volcan de Chiriqui, above Boquete, and about 
Cerro Punta. In eastern Chiriqui it has been collected at Cerro Flores, 
by Griscom, and on the Cordillera de Tolé (Salvin and Godman, Biol. 
Centr.-Amer., Aves, vol. 1 (pt. 13), 1881, p. 183). In Veraguas it was 
collected by Arcé at Calvovévora. 

The Collared Redstart behaves much like its congener, the Slate- 
throated Redstart, flying after insects and moving acrobatically through 


FAMILY PARULIDAE 21 


the trees or undergrowth. I have seen individuals of this species swing- 
ing on the side of a branch for an instant with wings and tail spread, 
so that they resemble a huge butterfly; many observers have remarked 
on their amazing tameness. At the elevations where both species of 
Myioborus occur they may both be found in the same mixed flock. On 
Cerro Punta, however, I have found torquatus at 1990 m in a humid 
cloud forest area where the trees were laden with moss and epiphytes, 
while miniatus occurred at the same elevation in a drier area. 

M. torquatus sings throughout the year, although not so frequently 
between July and February as does V/. miniatus. One song is an elabo- 
rate series of varied, mellow phrases, another is “a rather soft, slightly 
musical tszt-tsit-tsee, repeated” (Ridgely, 1976, p. 303). 

On February 26, 1955, I noted that all the Collared Redstarts on 
Cerro Punta were paired, but I have no information on the breeding of 
this species in Panama. Skutch, however, has observed several nesting 
pemsim Costa Rica (Pac..Coast. Avit., no. 31, 1954, pp. 371-376). 
The first nest he found was on April 3, when it was nearly completed. 
It and the other nests were placed on the ground, at the edge of a bank, 
in a hollow on a grassy hillside near woods, or beneath a decaying log. 
The nests were made of fine vegetable fibers, fibrous roots, bamboo 
leaves, and scales from tree ferns. Two or three eggs form the clutch; 
the eggs are white, sprinkled all over with light brown spots, most 
concentrated on the large end, and measure between 18.313.5 and 
19.1X13.5 mm. The incubation period is 15 days. 

In the few days just before and just after her eggs hatched, a female 
that Skutch watched would flutter helplessly on the ground in front of 
the nest. At hatching the young have pink skin and sparse natal down. 
By 10 days of age they are well feathered and at 13 days they leave the 
nest. Young hatched in April leave their parents by mid-June, when 
they are difficult to distinguish from adults. In August they seem to 
be paired. 


BASILEUTERUS TRISTRIATUS (Tschudi): Three-striped Warbler, 
Cerrojillo 


Myiodioctes tristriatus Tschudi, 1844, Archiv. f. Naturg., 10 (1), p. 283. (Valley 
of Vitoc, Junin, Peru.) 


Small; crown striped black and buff; upper surface yellowish olive; 
undersurface light greenish yellow. 

Description.—Length 113-123 mm. Adult (sexes alike), forehead 
and superciliary, extending to nape, buff; sides of crown black; central 
crown stripe olive-ocher, becoming buff on nape; rest of upper surface, 


314 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


including wing coverts, yellowish olive; remiges and rectrices dusky 
with outer webs edged yellowish olive; line through eye and auriculars 
largely black (except in tacarcunae); patches below eye and on side 
of neck buff; throat whitish; sides of breast, sides, and flanks light 
yellowish olive; rest of undersurface light yellow; underwing coverts 
white, tinged yellow. 

The Three-striped Warbler is a montane bird found from Costa Rica 
to northern Venezuela and northern Bolivia. Three races have been 
recognized in Panama, melanotis, chitrensis, and tacarcunae. Exami- 
nation of specimens of chitrensis, including the 14 taken by R. R. Ben- 
son at Chitra, Veraguas, from which the race was described by Gris- 
com (Amer. Mus. Novit., no. 280, 1927, p. 13) shows that it differs 
slightly in a series, but single individuals are not separable in some 
cases. Some specimens from Chitra show a somewhat darker chest, 
but lighter ones cannot be separated from melanotis. As there is no 
natural barrier between the birds of the Veraguas highlands and those 
of Chiriqui belonging to melanotis, the Veraguas population seems too 
poorly differentiated to merit subspecific status. 

This species is found in highland forest and woodland, where it 
moves through the lower level of trees, in thickets, and undergrowth, 
usually traveling in mixed species flocks. Its vocalizations include “a 
rushing series of chipping or ‘tsitting’ twitters composed of weakly 
clipping dry notes sprinkled at times with squeaky and harsh ones,” as 
well as single notes (Slud, Bull. Amer. Mus. Nat. Hist., vol. 128, 1964, 
p. 334). - 

The breeding behavior of this species has been investigated by Wil- 
liam H. Buskirk in Costa Rica. He found that the female constructs 
the nest and performs all the incubation herself. The male, however, 
participates in feeding the nestlings, and when the young are fledged 
each parent takes full responsibility for a single offspring. Ridgely 
(in litt.) transcribed the song of a bird at Cerro Jefe as “a very fast, 
high, jumbled tidedeteeadedecheeaweea’; he also heard the bird give 
single notes as a contact call. 


BASILEUTERUS TRISTRIATUS MELANOTIS Lawrence 


Basileuterus melanotis Lawrence, 1868, Ann. Lyc. Nat. Hist. New York, vol. 9, 
April, p. 95. (Cervantes, Costa Rica.) 

Basileuterus tristriatus chitrensis Griscom, 1927, Amer. Mus. Novit., no. 280, p. 
13. (Chitra, 4,000 ft. Pacific slope of Veraguas, Panama.) 


Characters.—Auriculars black; central crown stripe grayish buffy; 
broad superciliary stripe pale buffy gray. 


FAMILY PARULIDAE 315 


Measurements.—Males (9 from Chiriqui and Costa Rica), wing 
60.0-65.0 (62.6), tail 52.7-57.7 (55.1), culmen from base 10.3-12.1 
(16) tarsus 20.1-21:7-(20.9) mm. 

Females (7 from Chiriqui and Costa Rica), wing 56.0-59.0 (58.4), 
tail 48.5-54.2 (50.9), culmen from base 10.2-12.2 (11.7, average of 6), 
featens 20,2-22°2 (21.2) nim, 

Resident. Rare and local in the subtropical forests of Volcan de 
Chiriqui and adjacent Bocas del Toro, from 1080 to 2250 m, and in the 
mountains of Veraguas. Recorded in Chiriqui from above Palo Santo, 
west of El Volcan, above Boquete on Alto de Chiquero, and Bajo 
Mono, Chiriqui, and in Bocas del Toro from Camp Cylindro, at 1590 m 
on the Holcomb Trail (Blake, Fieldiana: Zool., vol. 36, no. 5, 1958, 
pp. 561-562). In Veraguas it is found lower, from 600 to 1200 m 
(Ridgely, 1976, p. 303); here it has been recorded from Chitra and 
Santa Fe, where Rex R. Benson collected 16 in 1925, and from the 
Cordillera del Chuct, where Arcé collected 2 in 1869. 

In February-March 1976 Ridgely (in litt.) found it to be common 
in forests above the Fortuna Dam site in central Chiriqui; there it was 
a conspicuous nuclear species of the flocks of the forest understory. 
The birds were found from 1050 m up. 


BASILEUTERUS TRISTRIATUS TACARCUNAE Chapman 
Basileuterus tacarcunae Chapman, 1924, Amer. Mus. Novit., no. 143, p. 6. (East 
slope, Mt. Tacarcuna, 4,600 ft., below Colombia-Panama line, Darién, Panama.) 

Characters.—Black absent from auriculars, replaced by light yel- 
lowish olive; central crown stipe more buffy yellow and orange in the 
center, with some feathers tipped yellowish olive. 

A male taken February 20, 1964, at Cerro Mali, Darién, had the iris 
dark brown, maxilla and tip of mandible fuscous; base of mandible 
dull brownish white; tarsus and toes verona brown; claws fuscous- 
black. 

Measurements.—Males (7 from Darién), wing 59.0-62.0 (61.3), 
tail 48.3-53.7 (52.2), culmen from base 11.0-12.3 (11.9), tarsus 19.7- 
2S (20.8), mim. 

Females (6 from Darién), wing 57.6-58.9 (58.2), tail 47.5-51.6 
(49:5), culmen from base 10.5-12.3 (11.4), tarsus 19.5-21.6 (20.1) 
mm. 

Resident. Rare and local in forest in the highlands of eastern 
Darién, where it has been collected on Cerro Tacarcuna and Cerro 
Mali. It has also been collected in Colombia, around the headwaters of 
the Rio Cuti, Choco Province. There are sight reports of what may be 


316 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


this race or an intermediate form from Cerro Jefe, eastern Province of 
Panama, where D. Hill and Ridgely found it several times between 900 
and 990 m (Ridgely, 1976, p. 303). 

In my few encounters with this species in Darién it has been in 
flocks moving through the undergrowth or the lower tree crown. 

This form has been given specific rank by some authors. It does not 
occur on the Cerro Pirre range, where its place may perhaps be taken 
by B. melanogenys ignotus. 


BASILEUTERUS CULICIVORUS GODMANI Berlepsch: Golden- 
crowned Warbler, Reinita Coronidorada 


Basileuterus godmani Berlepsch, 1888, Auk, 5, p. 450. (Chiriqui, Panama.) 


Small; crown striped with central stripe yellow-orange; upper sur- 
face grayish green; undersurface yellow. 

Description.—Length 115-123 mm. Adult (sexes alike), sides of 
crown to nane striped black; central area yellow-orange; superciliary 
and rest of central crown stripe light yellowish olive, extending to nape; 
dusky spot in front and behind eye; ear coverts olive-green, finely 
streaked with whitish; rest of upper surface dark greenish olive; wings 
grayish dusky with middle and greater coverts tipped light yellowish 
olive and outer web of remiges edged light gray; tail grayish dusky; 
sides of breast, sides, and flanks light yellowish olive; rest of undersur- 
face and bend of wing yellow; underwing coverts yellow and white. 

A male taken March 17, 1965, at El Volcan, Chiriqui, had the iris 
dark chestnut; lower half of rami and distal half of gonys pale brown- 
ish white; rest of bill fuscous-brown; tarsus and toes brownish yel- 
low; claws light brown; underside of toes yellow. 

Measurements.—Males (10 from Chiriqui), wing 57.4-63.0 (60.8), 
tail 48.7-55.5 (51.9), culmen from base 10.6-12.9 (11.9), tarsus 17.8- 
ZAM {OSB sasteale 

Females (10 from Chiriqui), wing 57.5-63.7 (59.9), tail 47.3-54.7 
(50.6), culmen from base 10.9-12.6 (11.8), tarsus 17.9-20.6+ (19m) 
mm. 

Resident. Fairly common in the lower highlands of Chiriqui and 
also known from the foothills of Veraguas and Herrera (Cerro Mon- 
tosa, 700-870 m) (Ridgely, 1976, p. 304). It is also found in Costa 
Rica. Other races are found in Mexico, through Middle America, and 
in South America east of the western Andes of Colombia to southern 
Brazil, Bolivia, Paraguay, and northern Argentina. Monniche col- 
lected this race on the Volcan de Chiriqui between 1710 and 1800 m 
(Blake, Fieldiana: Zool., vol. 36, no. 5, 1958, p. 562) and W. W. Brown, 


FAMILY PARULIDAE 327 


Jr., collected it there and at Boquete between 1200 and 2310 m (Bangs, 
Proc. New England Zool. Club, vol. 3, 1902, p. 60). Elsewhere in 
Chiriqui, it has been taken by Frank Hartman at Santa Clara (1230 m), 
by M. E. Davidson at Barriles (1350 m), and by Griscom at Cerro 
Flores (1200 m). Ridgely (im litt.) found it fairly common at I‘ortuna 
between 1000 and 1050 m in February-March 1976; it was replaced by 
B. tristriatus at higher elevations, but was not as common as that species. 
On the western slope of the Azuero Peninsula, Aldrich and Bole 
(Scient. Publ. Cleveland Mus. Nat. Hist., vol. 7, 1937, pp. 25-26) 
found this species uncommon in rain forest from 300 to 900 m and in 
cloud forest above 900 m. Arcé collected it in Veraguas at Calovévora 
(ealyin, Proc. Zool. Soc. London, 18/0, p. 183). 

The Golden-crowned Warbler inhabits forest and woodland, where 
it usually forages from the shrubbery up to about 7 m. It is often in 
pairs or little flocks, and regularly accompanies the small mixed flocks 
of the understory. At El Volcan, Chiriqui, on February 7, 1955, I saw 
a small scattered flock on a heavily wooded ridge of Cerro Picacho at 
1740 m. The birds moved with twitching tails through the smaller 
branches of the higher levels; in silhouette they were like the migrant 
warblers from the north. The call was a low chipping note. Slud (Bull. 
Amer. Mus. Nat. Hist. vol. 128, 1964, p. 334) describes the song as 
“three “wee’s on the same level, followed by three consecutively rising 
‘wee’s, with the last one accentuated. Usually, however, only the last 
three or so notes are given as an accelerating little crescendo or an un- 
musical “chip-chip-chirk’ lacking the ‘wee’ quality.” Buskirk et al. 
(Auk, 1972, p. 620) found that at Cerro Punta, Chiriqui, Golden- 
crowned Warblers regularly joined mixed flocks but showed only mod- 
erate tendencies to follow them. 

Nothing is known of this species’ breeding behavior in Panama, but 
Skutch (Publ. Nutt. Orn. Club., no. 7, 1967, pp. 150-154) has found 
a nest in Costa Rica, where this race also occurs. The nest was a globu- 
lar structure about 14 cm in diameter with a round opening on the side; 
it was set on the ground, hidden amid fallen leaves and twigs, in a 
fairly open forest. The outer surface of the nest was made of black 
rootlets, with additions of green moss, strips from the leaves of palms 
and branching stems of foliose liverworts; the thick inner cup was of 
very fine, brownish fibers. When Skutch found the nest on April 15, 
1964, it contained three eggs; they were white with a heavy wreath of 
dark brown blotches around the broadest part and a scattering of paler 
brown spots on the rest of the surface. Their measurements were 17.9 x 
14.1, 17.8X13.8, and 18.7 14.9 mm. 


318 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


On April 19 the young hatched; they had pink skin with sparse gray 
down. The young were fed mature and larval insects, and by their 
seventh day pin feathers had sprouted. These were open at 9 days, and 
the following day the young “exploded” from the nest when Skutch 
approached. At this time their feathers were still short, but the birds 
could fly a few feet and were led off by their parents. 


BASILEUTERUS MELANOGENYS Baird: Black-cheeked Warbler, 
Reinita Carisucia 

Small; crown striped chestnut and white; rest of upper surface gray- 
ish green; undersurface buffy white. 

Description.—Length 123-132 mm. Adult (sexes alike), forehead 
and very thin stripe on side of crown black; central crown area chest- 
nut, blackish at nape; rest of upper surface olive, becoming more 
greenish on rump; wings and tail dusky, with coverts tipped and remi- 
ges and rectrices edged yellowish olive; superciliary extending to nape 
white or pale yellowish; lores, orbital and auricular region black; sides 
of chin black, flecked white, central chin more white; throat whitish; 
broad breast band gray; sides, flanks, and abdomen olive; rest of under- 
surface buffy white; underwing coverts gray. 

This species is found only in the highlands of Costa Rica and Pan- 
ama, where in western Chiriqui it is common in forest, woodlands, and 
in brushy areas at the edge of pasture, while in Veraguas and in eastern 
Darién it is very local and only poorly known. | have found the Black- 
cheeked Warbler only in Chiriqui, where I have seen it in heavy forest 
and in brush along a small stream running through rough pastureland; 
it feeds in low undergrowth, often barely above the ground, up to mid- 
dle tree branches, usually moving in pairs. Skutch (Publ. Nutt. Orn. 
Club, no. 7, 1967, pp. 154-159) has studied this species in Costa Rica, 
and notes that it sings much less frequently than other species of Basil- 
euterus; he describes the song as slight and lisping. The bird also has 
weak chip call. 

Buskirk et al. (Auk, 1972, p.620) found that at Cerro Punta, Chiri- 
qui, the Black-cheeked Warbler was a short-term follower of the mixed 
species flocks that passed through its territory. Eisenmann (im litt.) 
only very occasionally saw this species with mixed species bands, and 
with these it usually foraged low; once only did he observe 1 well up in 
a tie: 
Three subspecies are currently recognized (all 3 found in Panama), 
nominate melanogenys, bensom, and ignotus. The last two were treated 
as full species by Todd (Proc. U. S.,Nat.;Mus)) vol:74, art; 19220noe 


FAMILY PARULIDAE 319 


79-80), although this treatment was not generally followed. The 
Check-list Committee of the American Ornithologists’ Union (Check- 
list of North American Birds, 6th ed., 1983, p. 627) has recently re- 
elevated ignotus to the rank of species, but there is no justification for 
this. 


BASILEUTERUS MELANOGENYS MELANOGENYS Baird 


Basileuterus melanogenys Baird, 1865, Rev. Amer. Birds, 1, p. 248. (Costa Rica.) 
Basileuterus melanogenys eximius Nelson, 1912. Smiths. Misc. Coll., 60(3), p. 22. 
(Boquete, 5,000 ft., Chiriqui, Panama.) 


Characters.—Superciliary white; side of head black. 

A male taken February 24, 1965, at Volcan de Chiriqui, Chiriqui, 
had the iris dark brown; cutting edge of maxilla and all of mandible 
flesh color; rest of maxilla fuscous-brown; tarsus and toes pale brown- 
ish white; claws dull neutral gray. 

Measurements.—Males (10 from Chiriqui), wing 56.5-66.5 (61.1), 
tail 58.1-62.3 (60.4), culmen from base 10.3-12.7 (11.7), tarsus 20.0- 
ZOO 2107))) ram. 

Females (10 from Chiriqui), wing 58.8-62.1 (60.3), tail 54.7-61.0 
(58.6), culmen from base 11.3-12.7 (11.9), tarsus 20.7-23.5 (21.9) 
mm. 

Resident. Common in forests, woodlands, and borders in the high- 
lands of western Chiriqui, and in adjacent Costa Rica. In Panama 
B. m. melanogenys has been collected at Cerro Punta, the Volcan de 
Chiriqui, and Boquete. Ridgely (1m litt.) has found it more numerous 
near Boquete, on the wetter side of the mountain, than above Cerro 
Punta. From February to May of 1901 W. W. Brown, Jr., collected 
11 specimens at Boquete and Volcan de Chiriqui between 1350 and 3060 
m (Bangs, Proc. New England Zool. Club, vol. 3, 1902, p. 60); Mon- 
niche found it at Volcan de Chiriqui between 1560 and 2750 m (Blake, 
imteldiana: Zool), vol. 36; no. 5; 1958, p: 562). 

The population of the Chiriqui highlands was originally described 
under the name eximius by E. W. Nelson (Smiths. Misc. Coll., vol. 60, 
no. 3, 1912, p. 22), but, in examination of a large series from Chiriqui, 
this population was found to be inseparable from nominate melano- 
Genny son Costa ixica,)\Mhere are, ditterences that appear to be due to 
specimen age—older specimens from Chiriqui are buffier below and less 
grayish on the back than fresher specimens and they thus resemble the 
Costa Rican birds in the Smithsonian series, all of which are older. 
Comparison of fresh material from both localities is desirable. 


320 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


Skutch (Publ. Nutt. Orn. Club, no. 7, 1967, pp. 154-159) has studied 
the nesting behavior of this species in Costa Rica, where on March 21, 
1963, he found a female building a domed nest on the ground in a steep 
ravine. The next day darkness and falling ash from an eruption of 
Volcan [raz 32 km away seemed to have caused the desertion of the 
nest. Skutch found another nest that year on June 3; this one was set 
in a nearly vertical mossy bank in a rather open forest with dense un- 
dergrowth of bamboo. The nest was oven shaped with a foundation of 
leaves and bamboo blades, sides and roof of bamboo leaves and pieces 
of fern fronds and roots, and lined with shredded vegetable fibers and 
brown scales from the fronds of large ferns. The nest was about 18 cm 
in height, 14 cm from side to side, and 12 cm from front to back. The 
interior measured about 6 by 5 cm in diameter by 7 cm in height. 

When Skutch found this nest it already contained two eggs. They 
were white, speckled with cinnamon-rufous, deeper and more heavily 
concentrated on the broader end. Both eggs measured 19.0 15.0 mm. 
The eggs hatched on June 10. The young had pink skin and long but 
sparse gray down. One of the young disappeared a few days after 
hatching, the other was beginning to develop feathers at 7 days, but 
had disappeared 4 days later when Skutch next visited the nest. Both 
before and after hatching of the eggs, the female had performed injury- 
feigning displays when Skutch approached the nest. 

On July 10, 1979, Eisenmann (im litt.) found individuals of this 
species at 1900 and 2030 m at Cerro Punta evidently carrying food to 
young; 1 with an insect in its bill persistently remained in the area 
where approached, chirping nervously. On September 20, 1959, he 
Saw pairs at several places in densely grown hillsides or in ravines near 
Cerro Punta, keeping very low near the ground. On February 24 and 
26, 1960, on the trail to Boquete, between 2330 and 2430 m several 
pairs were seen perched low in shrubbery at the forest edge. One pair 
was building a nest at the base of a tree on a slope beside the trail; 1 
bird carried material while the other accompanied, calling a sweet tseep. 


BASILEUTERUS MELANOGENYS BENSONI Griscom 

Basileuterus bensoni Griscom, 1927, Amer. Mus. Novit., no. 280, p. 12. (Chitra, 

4,700 feet, Pacific slope of Veraguas, Panama.) 

Characters.—Upper surface iron gray instead of greenish; under- 
surface white instead of yellow; with distinct gray pectoral band darker. 

Measurements.—Males (3 from Chitra, Veraguas), wing 57.5-63.0 
(60.2), tail 53.5-55.5 (54.6), culmen from base 11.4-11.8 (11.6), tar- 
sus 21.5-21.8 (21.6) mm. 


FAMILY PARULIDAE 321 


Females (3 from Chitra, Veraguas), wing 54.0-56.5 (55.3), tail 
51.5-52.2 (51.8), culmen from base 11.4-11.7 (11.6), tarsus 20.6-22.5 
ino!) tm. 

Resident. This race is known only from mountains about Chitra, 
eastern Veraguas, where it was first collected by Rex R. Benson in 
February 1926. He encountered it between 1200 and 1500 m, the latter 
altitude representing the tops of the highest peaks in that area. 

Eisenmann observes that in the Veraguas highlands several species 
develop forms with more black pigment in the plumage, while in Darien 
the tendency is to reduced melanin and more carotenoid yellow. 


BASILEUTERUS MELANOGENYS IGNOTUS Nelson 
Basileuterus melanogenys ignotus Nelson, 1912, Smiths. Misc. Coll., 60(3), p. 21. 


(Mount Pirri, 5,200 ft., near head of Rid Limon, Darién, Panama.) 

Characters.—Superciliary pale greenish yellow; undersurface with 
strong yellowish suffusion, black on head reduced. 

Measurements.—Male (1 from Cerro Tacarcuna, Darién), wing 
59.0, tail 57.0, culmen from base 12.8, tarsus 21.0 mm. 

emele (lirom Cerro Pirre, Darien, the type), wing 57-0, tail 55.4, 
culmen from base 11.5, tarsus 20.5 mm. 

Resident. Known only from eastern Darién, where the type, an 
adult female, was collected at Cerro Pirre (as 1560 m) near the head of 
the Rio Limon, on April 18, 1912, by FE. A. Goldman, and 3 males and 
2 females were collected by C. Myers and R. Hinds at 1650 m on Cerro 
Tacarcuna on February 9-12, 1975. Eisenmann, who examined all the 
Tacarcuna specimens in comparison with the type, noted that they 
showed much more black on sides of the head, forehead and throat 
than the lone Pirre specimen and a narrower superciliary. The Tacar- 
cuna birds thus seem nearer to the western Panama birds in this re- 
spect than to the type of ignotus. 


BASILEUTERUS RUFIFRONS (Swainson): Rufous-capped Warbler, 
Reinita Cabecicastana 


Setophaga rufifrons Swainson, 1838 (1837?), Anim. Menag., p. 294. (Real de 
Arriba, Mexico, Mexico.) 
Small; crown reddish brown; nape gray; rest of upper surface 
olive-green; undersurface yellow, greenish on sides. 
Description—Length 111-121 mm. Adult (sexes alike), super- 
ciliary white; lores, orbital ring, and line through eye black; crown 
and auriculars reddish brown; nape gray; rest of upper surface, in- 
cluding tail, olive-green; wings dusky with coverts tipped and remiges 


322 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


edged olive-green; chin, sides of throat, and side of face to auriculars 
white; sides of breast, sides, and flanks olive-yellow; rest of under- 
surface yellow. 

The Rufous-capped Warbler is found from northern Mexico 
through Central America to northern Colombia and western Venezuela. 
In Panama the race mesochrysus is common in lowlands of the Pacific 
slope and on the Caribbean slope in the vicinity of the Canal Zone. 
Another race, actuosus, inhabits Isla Coiba off the Pacific Coast of 
Veraguas. The forms ranging from southeastern Guatemala south- 
ward through Middle America to Colombia and Venezuela (including 
all Panama populations) are often separated as a distinct species, B. de- 
latri (see Todd, Proc. U.S. Nat. Mus. 74, art. 7, 1929, pp. 85-87; Hell- 
mayr, Field Mus. Nat. Hist! Zool., vel. 13, pt. 8; 1935) "pp. 508-912) 

The habitat of this species is scrubby clearings, weedy fields, and 
second growth. It usually forages near the ground, where its secretive 
behavior makes it hard to see, but once at Pesé, Herrera, I watched 1 
come up to feed actively in leafless branches 10 m above a stream. 
Skutch, who has observed this species in Costa Rica (Publ. Nutt. Orn. 
Club, no. 7, 1967, pp. 159-164) noted that it usually travels in pairs. 
Males sing from February to August and may deliver their song from 
elevated perches at dawn, but sing from within thick shrubbery during 
the rest of the day. Eisenmann (Smiths. Misc. Coll., vol. 117, no. 5, 
1952, p. 53) describes the song as heard at Playa Coronado in June, as 
“a rather rapid, dry, chittering chit-cha-chup-cha-chuweép, with vari- 
ations. ... calls include a simple chit, a zeep-zeep or dzit-dzit.”” Some- 
times the song ends witha chee-weécha. In central Panama, Eisenmann 
has noted singing from early March through late June. 

E. A. Goldman collected 2 specimens with full stomachs at Corozal 
in the Canal Zone on June 15, 1911: one contained a chrysomelid 3%, 
a small curculionid 2%, elaterid fragments 3%, bug remains 3%, many 
fulgorid remains finely ground 44%, more than four large ants 35%, 2 
smaller species 8%, other coleoptera 2%; the other contained 5 ants 
44%, a reduviid 15%, Achalles sp. 10%, 2 cerambycids 15%, a riti- 
dulid near Colastus 8%, other coleoptera fragments (Chrysomelidae) 


Sia 


BASILEUTERUS RUFIFRONS MESOCHRYSUS Sclater 
Basileuterus mesochrysus P.L. Sclater, 1861, Proc. Zool. Soc. London, 28 (1860), 
p. 251. (“Bogota,” Colombia.) 
Characters.—Undersurface brighter yellow than in B. r. actuosus; 
gray of nape lighter. 


FAMILY PARULIDAE 323 


A female collected February 21, 1962, at El Copé, Coclé, had the 
iris dark mouse brown; bill black; tarsus, toes, and claws light brown 
with a line on back of tarsus buffy brown; pads honey yellow. 

Measurements.—Males (10 from Panama), wing 50.3-56.9 (54.3), 
tail 47.1-55.0 (50.0), culmen from base 11.2-13.5 (12.2), tarsus 19.3- 
20.6 (20.0) mm. 

Females (10 from Panama), wing 50.2-57.8 (53.8), tail 46.5-53.7 
(49.5), culmen from base 11.5-13.8 (12.4), tarsus 18.8-20.6 (19.6) 
mm. 

Resident. Fairly common throughout the mainland Pacific slope 
lowlands from Chiriqui to the Canal Zone area and on the Caribbean 
slope in the vicinity of the Canal Zone. In Chiriqui it has been col- 
lected as high as 1560 m (Blake, Fieldiana: Zool., vol. 36, no. 5, 1958, 
p. 562), and in central Panama it has been recorded to 950 m on Cerro 
Azul. Aldrich and Bole (Scient. Publ. Cleveland Mus. Nat. Hist., vol. 
7, 1937, p. 23) found it uncommon in brushy savannas and forest 
margins near Montijo Bay, Veraguas. In 1911, E. A. Goldman re- 
corded it as “‘a rather common bird in the partly open country through- 
out the Zone.” This race also occurs in southwestern Costa Rica and 
in northern Colombia and western Venezuela. Griscom (Bull. Mus. 
Comp. Zool, vol. 78, 1935, p. 370) states it ranges to Darién, but I am 
not aware of any specimens or sight reports from that province. 

According to Eisenmann, this bird favors thickets or lightly-wooded 
semi-arid areas, but in more humid habitats follows clearings into 
second growth and woodland borders up into the lower highlands and 
to the Caribbean coast in the Canal Zone and adjacent parts of Colon 
Province. If forest grows up again in humid areas it tends to disap- 
pear, as on Barro Colorado Island (Willis and Eisenmann, Smiths. 
Contrib. Zool. 291, 1979, p. 27). 

A bird banded by H. Loftin (im litt. to Eisenmann) at Curundu, 
Canal Zone, on January 8, 1963, was recaptured there on March 31, 
1966. 

Peas Goldman shota pair at Gatun, Canal Zone, on’ May 3, 1911; 
that were in breeding condition; the female contained two large eggs, 
the larger of which was nearly ready to lay. A pair that I collected 
March 7, 1960, at Buena Vista, Chiriqui, was also in breeding con- 
dition. Kisenmann noted a fledgling being fed in the Canal Zone on 
June 25. 

Skutch (Publ. Nutt. Orn. Club. no. 7, 1967, pp. 159-164) has found 
the nest of this species in Costa Rica. Like other members of the genus, 
B. rufifrons places the nest on the ground where it is hidden amid litter 


324 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


or set between rocks or against a fallen log. The nest is oven shaped 
and constructed of fine herbaceous stems, grass blades, fragments of 
dead leaves, rootlets, and other fine plant material. The bottom is a 
thick pad of finely shredded bast fibers. One nest measured 12 cm high, 
12 cm from back to front, and 14 cm from side to side. The inside 
measurements were 6 cm high, 11 cm from front to back, and 7 cm from 
side to side. 

All the nests in which Skutch found eggs contained a clutch of three. 
Those from one set were white, marked with fine specks of cinnamon 
crowded over the thicker end and more sparingly scattered over the rest 
of the surface. They measured 17.9 14.3, 17.5x14.3, and 17.5x14.3 
mm. Incubation is performed only by the female; Skutch could not 
determine the length of the incubation period. The young are born 
with yellow mouth cavities and sparse gray down on pink skin. They 
are fed by both parents, who bring them insects including long green 
caterpillars. In one nest the feathers of the young were expanding 
rapidly at 8 days of age and at 12 days, when their upper surface was a 
uniform dark gray without head markings and their undersurface yel- 
lowish olive, the young left the nest. 

Two females collected in Panama by Strauch (Bull. Brit. Orn. Club, 
1977, p. 64) weighed 10.1 and 11.8 g. 


BASILEUTERUS RUFIFRONS ACTUOSUS Wetmore 
Basileuterus delatru actuosus Wetmore, 1957, Smiths. Misc. Coll., 134 (9), p. 92. 

(Isla Coiba, Panama.) 

Characters.—Bill larger and coloration darker than that of B. 1. 
mesochrysus. | 

M casurements.—Males (11 from Isla Coiba), wing 57.6-62.5 (60.1), 
tail 51.1-55.8 (53.3), culmen from base 13.3-14.4 (13.8), tarsus 20.5- 
Pate (As) \) agua, 

Females (5 from Isla Coiba), wing 56.1-60.2 (57.5), tail 50.0-53.5 
(51.8), culmen from base 13.5-13.9 (13.6), tarsus 20.0-21.6 (21.0) 
mm. 

Resident. Common on Isla Coiba, off the Pacific coast of Veraguas. 
In January 1956 I encountered birds near the shoreline and back 
through high forest in the interior of the island. They also came into 
thickets in abandoned fields near the work camps. They were common, 
but of secretive habit, keeping behind cover. As they frequently carry 
their tail at an angle over the back, they often suggest wrens as they 
move about behind the screening twigs and leaves. At this season they 
were silent except for an occasional chipping call. Usually they are 


FAMILY PARULIDAE 325 


found in pairs, and during January most of the specimens I obtained 
were nearly ready to breed. 

Although the total length of the bill in the Coiba Island birds is only 
slightly more than in mainland individuals, the breadth and general 
bulk are appreciably greater. The darker color of this race is a general 
tendency in the races endemic to Coiba. 

On April 13, 1976, Ridgely (in litt.) saw a pair on Coiba with two 
fledged but still dependent young. He was interested to note that the 
birds were common in forest undergrowth, a habitat in which they 
would never be found in on the mainland. 


BASILEUTERUS FULVICAUDA (Spix): Buff-rumped Warbler, 
Reinita de Rabadilla Anteada 


FIGuRE 25 


Muscicapa fulvicauda Spix, 1825, Av. Spec. Nov. Brasil, 2, p. 20, pl. 28, fig. 2. 
(Sao Paulo de Olivenga, Rio SolimGes, Brazil.) 


Small; upper surface dark olive; upper tail coverts and basal half of 
rectrices buffy; outer half dark brown; undersurface buffy, marked 
with dark olive. 

Description.—Length 118-128 mm. Adult (sexes alike), crown 
blackish olive, becoming dark olive on rest of upper surface to upper 
tail coverts, which are warm buff; wing coverts and remiges blackish, 
edged dark olive; basal half of tail ochraceus-buff, outer half dark 
brown; fine superciliary warm buff; undersurface white mottled with 
varying degrees of warm buff and dark olive. 

Immature, like adult, but throat and breast heavily marked with dark 
olive. 

There is little justification for separating this species and its ally B. 
rivularis in the genus Phaeothlypis as was done by Lowery and Monroe 
(in Peters, Check-list Birds World, vol. 14, 1968, p. 75). Todd (Proc. 
U.S. Nat. Mus., no. 2752, 1929, p. 8) erected the genus Phaeothly pis 
only for the forms here included in the species fulvicauda. The char- 
acters he used to distinguish the genus were “bill relatively wider, . . 
tail relatively shorter, much less than the distance from the bend of the 
wing to the end of the longest secondaries, . . . style of coloration very 
different, the tail being always bicolor...’ Hellmayr (Field Mus. Nat. 
Hist. Zool., Cat. Birds. Amer. part 8, 1935, p. 522) did not recognize 
Phaeothlypis for fulvicauda on account of the recognizably close re- 
lationship of that species with rivularis, a species that Todd included in 
Basileuterus. None of Todd’s characters for Phaecothly pis apply to B. 
rivularis, in which the tail is longer and not bicolored, and in some 


326 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


forms of which, such as bolivianus, the bill is actually narrower than 
in a number of more distantly related species of Basileuterus (e.g., 
coronatus). Lowery and Monroe, by including rivularis in Phaeo- 
thly pis, left the genus with no distinguishing characters. The best way 
to resolve this inconsistency is by refusing to recognize the validity of 
Phaeothly pis. 


Froure 25.—Buff-rumped Warbler, Reinita de Rabadilla Anteada, Basileuterus 
fulvicauda. 


In Panama there are two very distinct forms of B. fulvicauda that 
Todd (op. cit., p. 9) considered to be separate species. One form, 
semicervina, is found in the eastern half of Panama and in the Western 
and Central Andes from Colombia to northwestern Peru. Two sub- 
species have traditionally been recognized within the western half of 
Panama, leucopygia and veraguensis (with their respective synonyms 
gaffneyi and toddi). The stated range of leucopygia is from Honduras 
through Costa Rica and along the Caribbean slope of Panama to Ver- 
aguas; that of veraguensis is from southwestern Costa Rica along the 
Pacific slope of Panama to the Canal Zone. According to Todd (p. 
12), the only distinguishing feature of veraguensis is the supposedly 
buffier, less whitish, underparts. A series of 36 specimens from Nica- 
ragua to the Canal Zone shows individual variation to be too great to 
permit any subdivision of this population. Three birds (USNM no. 


FAMILY PARULIDAE 327 


469102-4) taken in 1958 on the Western River, Almirante, Bocas del 
Toro, show this variability admirably. One is light underneath, as at- 
tributed to typical leucopygia, another is much buffier, while the third 
is extremely dark, the underparts except the throat and midline of 
belly being entirely suffused with dusky. Certain individuals from 
Chiriqui are as light as any from farther north, while others from 
Nicaragua and Costa Rica are as buffy as some from Coclé or Chiriqui. 
Therefore, these populations are listed under the name leucopygia, 
with veraguensis as a junior Synonym. 

There is a narrow zone of intergradation between leucopygia (sensu 
lato) and semicervina in the Canal Zone and slightly eastward. Two 
birds collected by E. A. Goldman labeled “Rio Indio, Canal Zone” (a 
tributary of the Chagres, not the following Rio Indio in western Colon 
Province) and another taken from Chilar, on the Rio Indio, Colon, are 
typical of leucopygia, as are all those from farther west. Birds from 
Cerro Chucanti, Serrania de Majé, eastern Panama Province, and from 
there east through San Blas and Darién and most of Colombia are 
typical of semicervina. Except for the 2 Goldman specimens above, 
all birds from the Canal Zone proper (Gamboa, Gatun, Cocoli River), 
Cerro Azul, Utivé, and the upper end of Madden Lake in eastern 
Province of Panama (Candelaria and Peluca Hydrographic Stations) 
are clearly intergrades. An exception is an individual from the Peluca 
Station (1 of 4 taken there) that appears perfectly typical of semicer- 
vina, suggesting that the zone of intergradation probably does not ex- 
tend much farther east than this locality. 

Some authors merge the B. fulvicauda complex in B. rivularis 
(Riverside Warbler) of eastern, Amazonian, and southern South 
America south to northern Argentina (Meyer de Schauensee, Species 
Birds S. Amer., 1966, p. 453). 

The Buff-rumped Warbler dwells along the shores of streams or, 
more rarely, boggy areas and mangrove swamps. Eisenmann has gen- 
erally seen this species only at rapid-flowing, narrow, clear streams over 
rocky beds, but in coastal Bocas del Toro east of Almirante, on the 
slow-moving, dark, muddy, rather wide Western River, on October 19, 
1965, he saw two pairs at different places in waters navigable by motor 
launch. In many aspects of behavior this species resembles a water- 
thrush, but it hops rather than walks, fanning and wagging its tail 
from side to side, exposing the conspicuous light patch on the upper 
tail coverts and base of the tail feathers. I have always found this a 
rather shy species, continually flying ahead and rarely affording one a 
good look. 


328 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


The song is varied and clear, with final phrases like the loud trilling 
of a domestic canary. Skutch (Pac. Coast Avif., no. 31, 1954, p. 348) 
found that the female sings a rich warble in response to the song of the 
male. This species sings more or less throughout the year, but most 
during the nesting season; the song is usually delivered from the ground 
or a rock or log in a stream. Eisenmann describes the song as explo- 
sive, emphatic, and loud (usually increasing toward the end), with 
notes in rapid succession often given from a perch, 5-3 m above the 
ground, above or near a stream. On April 12 and 16, 1959, he wrote 
songs as chewy, chewy, chee-chee-chee-chee-chee-chee-, choo, choo, 
choo, choo-choo-choo-choo, tyoo, tyoo. The song lasted 4-6 seconds; 
the numbers of chee and other notes varied, and the tyoo notes might be 
omitted. The call is a loud, sharp note or a series of such notes. 

The stomach of | collected by E. A. Goldman at Cana, Darien, con- 
tained bits of two dragonflies 20%, fragments of an ant 10%, an 
ichneumon 5%, dipteran remains finely ground 54%, fragments of a 
cerambycid 5%, bits of two other beetles 6%. 


BASILEUTERUS FULVICAUDA LEUCOPYGIUS Sclater and Salvin 


Basileuterus leucopygius P.L. Sclater and Salvin, 1873, Nomen. Av. Neotrop., 
pp, 10) 156) (Costa Ricaz) 

Basileuterus [leucopygius subsp. a] veraguensis Sharpe, 1885, Cat. Birds. Brit. 
Mus., 10, p. 403. (Paraiso Station, Panama.) 

Basileuterus fulvicauda gaffneyi Griscom, 1927, Amer. Mus. Novit., no. 280, p. 14. 
(Guaval, Rio Calovévora, ... Veraguas, Caribbean slope, western Panama.) 
Basileuterus fulvicauda toddi Griscom, 1927, Amer. Mus. Novit., no. 280, p. 14. 

(Boqueron, Chiriqui, western Panama.) 


Characters.—Undersurface mottled and smudged with dark olive; 
upper surface and cheeks darker; superciliary obscure; dark areas of 
tail, particularly the outer rectrix, more extensive. 

A male collected at Puerto Armuelles, Chiriqui, March 8, 1966, had 
the iris dark brown; base of mandible very dull dark brown; rest of bill 
black; inside of mouth pale bluish gray; tarsus and toes brown; claws 
dull slate. 

Measurements.—Males (10 from western Panama), wing 61.0-66.9 
(64.4), tail 50.0-55.7 (52.4), culmen from base 12.7-14.9 (14.1), tarsus 
Z13-22.9. (22:2) ama: 

Females (10 from western Panama and Costa Rica), wing 60.4-65.5 
(62.7), tail 47.1-54.6 (51.2), culmen from base 12.8-14.5 (13.8), tar- 
sus 19.0-22.6 (21.3) mm. 

Resident. Uncommon to fairly common along streams and rivers 


FAMILY PARULIDAE 329 


in forest and second-growth woodland of both slopes from the Costa 
Rican border to central Panama, where it is less frequent and inter- 
grades with the race semicervinus, as described above. It has been 
found as high as 1620 m, where Monniche collected it at Horqueta, 
Chiriqui (Blake, Fieldiana: Zool., vol. 36, no. 5, 1958, p. 563). Ridgely 
(in litt.) finds that during the dry season it seems to disappear from 
areas of the Canal Zone where streams mostly dry up, e. g., around 
Summit and in Madden Forest. 

Skutch (Pac. Coast Avif., no. 31, 1954, pp. 346-356) found that in 
Costa Rica the nesting season runs from March to August. Both sexes 
build the nest, which is oven shaped and usually placed in a niche on a 
steep slope, often close to a stream or river. The nest takes up to 13 
days to construct. It is made of coarse rootlets, dry grass blades, leaf 
fragments, and weed stems; the lining is of fine shredded fibers and 
leaf skeletons. Two eggs form a clutch; they are glossy white, marked 
around the blunt end with a heavy wreath of brown blotches or spots 
and thinly spotted with the same shades over the remaining surface. 
The average measurements of twenty eggs are 20.7X14.9 mm. In- 
cubation is performed entirely by the female and requires 16 to 19 
days. When the young hatch they have pink skin and scanty gray 
down. The interior of the mouth is yellow. They are fed by both par- 
ents and usually remain in the nest until 13 or 14 days old, when they 
are well feathered. Only a single brood is reared each year. 


BASILEUTERUS FULVICAUDA SEMICERVINUS Sclater 


Basileuterus semicervinus P.L. Sclater, 1861, Proc. Zool. Soc. London, 28 (1860), 
p. 84. (Nanegal, Ecuador.) 


Characters.—Breast unmarked; cheeks buffy; buffy superciliary 
more prominent; upper surface lighter; light area of tail more ex- 
tensive. 

A female collected at Armila, San Blas, on February 23, 1963, had 
the iris dark brown, base of gonys and lower half of mandibular rami 
dull brownish white; rest of bill black; tarsus and toes brownish honey 
yellow; claws light mouse brown. 

Measurements.—Males (10 from eastern Panama), wing 61.5-66.0 
(63.6), tail 49.0-54.8 (51.7), culmen from base 12.7-15.3 (13.8), tar- 
sus 19.9-22.3 (21.4) mm. 

Females (10 from eastern Panama), wing 58.9-64.0 (60.6), tail 45.8- 
92.5 (49.7), culmen from base 12.3-14.8 (13.5), tarsus 19.3-22.6 
(211.2) mar. 


330 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Resident. Uncommon to fairly common in the lowlands and foot- 
hills of both slopes from east of the Canal Zone to the Colombian 
border. I have found it as high as 1260 m, on the south fork of the Rio 
Pucro) Cerro MahDanicn: 


ZELEDONIA CORONATA Ridgway: Zeledonia, Zeledonia 
FIGURE 26 


Zeledonia coronata Ridgway, 1889, Proc. U.S. Nat. Mus., 11 (1888), p. 538. 
(Laguna del Volcan de Poas, Costa Rica.) 


Small; center of crown orangey brown, margined by black; rest of 
upper surface brownish olive; forehead, sides of head, and undersur- 
face slaty. | 

Description.—Length 100-110 mm. Adult (sexes alike), center of 
crown orangey brown, bordered by narrow black stripe; forehead and 
side of head dark slate gray; rest of upper surface brownish olive; 
undersurface slaty gray; flanks and undertail coverts dull olive; 
remiges and rectrices dark brown, edged brownish olive. 

Juvenile, like adult, but lacks crown patch; underparts olive. 

Measurements.—Males (10 from Veraguas, Chiriqui, and Costa 
Rica), wing 61.5-69.5 (65.8), tail 36.3-42.3 (38.8), culmen from base 
11.7-13.5 (12.8, average of 9), tarsus 26.1-30.9 (28.7) mm. 

Females (10 from Veraguas and Costa Rica), wing 61.0-68.0 (62.9), 
tail 33.3-38.8 (35.8), culmen from base 11.8-13.9 (12.7), tarsus 26.2- 
29.50 (2727) mama 

Resident. Rare and local in the highlands of Chiriqui and Veraguas, 
where it inhabits very humid forests with thick ground cover. It has 
been recorded in Chiriqui between 1740 and 3090 m and in Veraguas at 
Chitra from 1050 to 1200 m (Ridgely, 1976, p. 305). In Chiriqui it 
has been collected on the Volcan de Chiriqui and at Boquete. Ridgely 
heard and briefly saw one in cloud forest well above Fortuna (1800 m) 
in central Chiriqui on February 27, 1976, and N. G. Smith reports (1 
itt. to Eisenmann) seeing one on Cerro Colorado, eastern Chiriqui, in 
April, 1979. The Zeledonia is also found in Costa Rica. 

The systematic position of Zeledonia coronata was formerly the sub- 
ject of considerable uncertainty. It was thought, on quite inadequate 
grounds, to be related to thrushes, although equivocation resulted in 
its being placed in a monotypic family Zeledoniidae. Sibley (Postilla, 
no. 125, 1968, pp. 1-12), after a review of its taxonomic history and an 
analysis of the electrophoretic pattern of its egg-white proteins, con- 
cluded that this species is not a thrush, but probably a parulid, on the 


FAMILY PARULIDAE Gia 


basis of egg-white data and morphology. The parulid affinities of Zele- 
donia were confirmed by Raikow’s (Bull. Carnegie Mus. Nat. Hist., 7, 
1978, p. 28) dissections of the musculature, which also suggest that 
Zeledonia may be more nearly related to Basileuterus. As the name 
“Wrenthrush,” formerly applied to this bird, is now seen to be inap- 
propriate, the name “Zeledonia” has been used in its place. 


x. TF 


FicurE 26.—Zeledonia, Zeledonia, Zeledonia coronata. 


Until James Hunt studied it in Costa Rica during 1968 (Auk, 1971, 
pp. 1-20), little was known of the biology of Zeledonia except its 
whistled call and that it was a very secretive inhabitant of dense under- 
growth in the humid herb layer of high mountains and in tangles and 
vines on trunks and lower limbs directly above it, where it hops and 
flicks its wings, but very rarely flies. The Zeledonia’s commonest call 
is a “thin, high-pitched whistle, usually with a slight rising inflection,” 
transcribed by Ridgely as a very piercing, sibilant pseee. The song, 
which Hunt first heard on March 19, is a sequence of phrases like 
sseee-del-deet with emphasis on the last syllables. 

Hunt found a nest under construction on March 25. It was situated 
in a cavity in a vertical mossy bank of a small stream. The nest, finished 
on March 31, was a domed structure with an entrance on the side, 
made of mosses with a few small leaves and twigs and lined with fine 
dead plant material, including grass, mosses, and decaying leaves. The 
first egg was laid on April 8. It and a second, laid 2 days later, were 


2e0 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


white or buffy white marked with brown spots. They measured 21.9X 
17.3 and 21.2X16.9 mm. 

This nest was abandoned, but another found May 21 was identical 
in appearance and situation. It contained 2 young birds probably not 
more than 3 days old. They were sparsely covered with down and their 
eyes were still shut. Their mouth linings were orange. The young of 
this nest died a few days later, but a third nest found June 26 had 2 
nestlings about 10 days old. By 15 days these young were fully 
feathered; 2 days later, when Hunt had to leave the area, the nestling 
he had not collected was still in the nest, indicating a longer pre-fledging 
period than that of any other wood warbler. 


Family ICTERIDAE: Blackbirds and Orioles, Turpiales y Chacareros 


The Icteridae are an exclusively American family that has its great- 
est diversity in the tropics. Twenty-two species have been found in 
Panama, including 5 migrants from temperate North America. The 
icterids are medium- to large-sized birds, mostly boldly patterned in 
black with yellow, red, or orange, or entirely black. The oropendolas 
are the largest passerines in the tropics; along with the caciques and the 
Giant Cowbird, they have the ability to produce loud noises with their 
wings when in flight that are sometimes part of courtship displays. An- 
other peculiarity of many of the larger icterids is a strong musky odor. 

The icterids are particularly interesting for the diversity of their 
breeding behavior. The oropendolas and some of the caciques live in 
colonies in isolated or exposed trees, where females outnumber males 
and weave long, pendulous nests. Other caciques and the orioles nest 
in solitary pairs, while the Red-breasted Blackbird and Eastern Mea- 
dowlark sometimes form loose colonies in large fields and savannas. 
The Panama cowbirds are parasites with varying degrees of special- 
izations, from the Shiny and Bronzed Cowbirds, which put their eggs 
in the nest of a variety of small birds, to the Giant Cowbird, which 
parasitizes only the colonial oropendolas and caciques. 

All the icterids have powerful voices, some of which are loud and 
raucous, while others produce very fine songs. Their diet is varied, in- 
cluding insects, fruit, seeds, and nectar. The Great-tailed Grackle regu- 
larly scavenges and takes small vertebrate prey including nestling birds; 
there are records of oropendolas snatching birds as well. 

Although the Dickcissel, Spiza americana, is usually included with 
one or another of the finchlike groups, Raikow’s (Bull. Carnegie 
Mus. Nat. Hist., vol. 7, 1978, pp. 1-43) examination of the limb myol- 


FAMILY ICTERIDAE 333 


ogy corroborates the observations of Beecher (Auk, 1951, pp. 411- 
440) based on jaw muscles and shows S‘piza to be a primitive member 
of the Icteridae, close to the Emberizinae. This conclusion is followed 
here, aS no evidence in support of a contrary position has been 


presented. 
KEY TO SPECIES OF ICTERIDAE 
_ PEPRALISIG7 DIZ YOKE ACES INE DES OL a ee ane IAAT Artin) cis RE MR a eo je 
‘hsih Bimal hig Td eye) cca ae eaten alacali Me Aa RG Mae Uae SET te nnt A A a a 6 
SP Nea UCR Le Se yh a Na Lc ay tart. ca le A My CMU ARE a MU OAV SALE a 3 
ILE ERIT-GUZM AURATS A aie A Fe Rn Ou NUT cCIC TY 7. TR a See a 4 
3. Tail not graduated. 


10. 


lid 


12. 


£3. 


Giant Cowbird, Scaphidura oryztvora. p. 355 
Tail graduated. 
male Great-tailed Grackle, Quiscalus mexicanus peruvianus. p. 362 


. Strong purple sheen. 


male Shiny Cowbird, Molothrus bonariensis cabanisu. p. 361 
ipleronine iiiaescent Sheen: oc Uik 1. cee UIE RN eel 5 


. Long light-colored bill. 


Yellow-billed Cacique, Cacicus holosericeus holosericeus. p. 353 
Short dark bill. 
Bronzed Cowbird, Molothrus aeneus aeneus. p. 358 


nO sa edit y KOT NG INOW hte eh elle eps ie oe ot ak ved We Misiaw le she dle Ola dlalace-eve « 7 


Rea KeduomMmoldhy Patctermed ic ai.cketeiese ices omeerneees <lole eaters SN RINE is. 5 8 


. Large, with long graduated tail. 


female Great-tailed Grackle, Quiscalus mexicanus peruvianus. p. 362 
Smaller, tail not graduated. 
female Shiny Cowbird, Molothrus bonariensis cabanisu. p. 361 


. Large, body black or dark brown, tail yellow with central pair of rec- 


VESPLGEIS- SUG RI repeal er OPER TL GN BRE A 9 
Mail notall yellow, except for central pair of rectrices. 0500. Pee 12 


. Remiges and entire back reddish brown. 


Montezuma Oropendola, Psarocolius montezuma. p. 336 
Re eSM ACK Ot DIAGKASIT ey si ecre cottey oi rliacn te late van ney cy Sina Neg fori eiaeea ee 10 
Back and wing coverts brown. 
Black Oropendola, Psarocolius guatimozinus, p. 339 
Minnoce entirely mplack Om blackish. ee dees eens Cece a et Seid sierets e 3.e 11 
Head dark brown. 
Chestnut-headed Oropendola, Psarocolius waglert. p. 341 
Head black or blackish. 
Crested Oropendola, Psarocolius decumanus melanterus. p. 344 
Rump uniform with back and tail; head and/or chest orange-yellow. 
Yellow-headed Blackbird, Xanthocephalus xanthocephalus. p. 377 
RAamMpNCIntehs, [LOM COlOL OL Dalek Of tally. seek oe ee es ke ws 13 
Rump red. 
Scarlet-rumped Cacique, Cacicus uropygialis. p. 349 
Rep OLATe Citys eatay Sy Mntwie ye iat git Nak, lea mame New iains uRO udi Niki eh 14 


334 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


14. Broad buffy band across nape, rump white. 
male Bobolink, Dolichonyx oryzivorus 

in alternate (breeding) plumage. p. 381 
Rump. notswihites esheets Mae gee Se 15 

15. Undertail coverts yellow, rest of undersurface black. 
Yellow-rumped Cacique, Cacicus cela vitellinus. p. 346 
Most of undersurface ‘not ‘black... 2.0. 20%. 12... 344-0 eee 16 
16. Throat to belly bright red, rest of body black, or body streaked brown with 

patch of red on belly and bend of wing. 

Red-breasted Blackbird, Letstes militaris militaris. p. 378 


Novred on. body. scat cas Gone w onic ccs Raikes 17 
17. Upper surface streaked: brown. .. 0.2. ..4 2... 5-6 5 -k eee 18 
Upper surface not streaked. o.c65 co ec ed oom ee eee ee 21 
18. Undersurface yellow, with black patch. ....-...-. ...3¢6se=eeeenee 19 
Undersurface buffy seo!) 5 keel Bede s 04 anh eee Sr 20 


19. Large, more than 180 mm. 
Eastern Meadowlark, Sturnella magna subulata. p. 379 
Small, less than 155 mm. 
adult male Dickcissel, Spiza americana. p. 334 
20. Crown with broad, buffy central stripe. 
male in basic plumage or female Bobolink, Dolichonyx oryzivorus. p. 381 
Crown without broad stripes. 
female or immature Dickcissel, Spiza americana. p. 334 
21. Undersurface rich brown or yellowish green. 
Orchard Oriole, Icterus spurius spurius. p. 365 
Undersurface yellow or orange...) 2... Je yseese) eee ee 22 
22. Undersurface bright orange or washed with orange. 
Northern Oriole, Icterus galbula galbula. p. 375 
Undersurface yellow and black... «2.5 :.<...2.052. 299s Joe ee ee 23 
23. Entire back yellow. 
Yellow-backed Oriole, Icterus chrysater giraudu. p. 373 


24. Head black. 
Black-cowled Oriole, Icterus dominicensis praecox. p. 367 
Heads yvellowvor oranges)... 62d idos dud nardeval steely 2. eee 25 
25. Head orange. 
Orange-crowned Oriole, [cterus auricapillus. p. 368 
Head yellow. 
Yellow-tailed Oriole, Icterus mesomelas. p. 369 


SPIZA AMERICANA (Gmelin): Dickcissel, Arrocero 
Pasajero Americano 


Emberiza americana Gmelin, 1789, Syst. Nat., 1(2), p. 872. (New York.) 


Medium size; upper surface mostly streaked brown; undersurface 
white with yellow wash on breast; male with black lower throat. 

Description —Length 135-154 mm. Adult male in alternate plum- 
age, lores white, superciliary rich yellow; fine line around lower rim 


FAMILY ICTERIDAE 335 


of eye white; crown, nape, sides of head and of breast gray; crown 
tinged greenish; rest of upper surface brown, with upper back streaked 
blackish; wing coverts reddish brown; remiges and rectrices blackish 
brown; tertials edged light brown; upper throat white; moustachial 
stripe begins white, becomes rich yellow and terminates white; lower 
throat and upper breast black in shape of a V; rest of breast and center 
of belly rich yellow; rest of undersurface white; bend of wing dark 
yellow; underwing coverts white, tinged yellow. 

Male in basic plumage duller, with feathers of black bib tipped buff. 

Female, like male, but duller, with crown and nape light brown in- 
stead of gray, a thin gray stripe on sides of throat and across bottom 
of throat; yellow less extensive and black absent. 

Measurements.—Males (10 from the breeding range, taken in May), 
wing 78.1-84.8 (82.1), tail 51.4-61.5 (58.0), culmen from base 14.3- 
15.8 (15.1), tarsus 21.4-23.4 (22.4) mm. 

Females (10 from the breeding range, taken in April and May), 
wing 70.5-77.2 (74.6), tail 50.3-58.6 (54.3), culmen from base 13.3- 
15.5 (14.6), tarsus 19.3-22.6 (21.3) mm. 

Migrant and winter visitor from the north. Fairly common and 
sometimes abundant as a migrant on the Pacific slope; on the Carib- 
bean slope more erratic. Less common in winter, mainly on the Pacific 
slope. Most of the population winters in northern South America. In 
Chiriqui, the Dickcissel has been collected as high as 1200 m at Boquete, 
where W. W. Brown, Jr., took a male on January 24, 1901 (Bangs, 
Proc. New England Zool. Club, vol. 3, 1902, p. 70). In January of 
1956, I found 2 in a pasture on Isla Coiba, off the Pacific coast. This 
species is commonest during September and April, but first appears in 
late August (Ridgely, 1976, p. 332) and stragglers remain until May 
and June—there is 1 specimen from June 16 (Eisenmann, Smiths. 
MiserColls vol 117, no. 5, 1952, p. 57). One banded at Almirante, 
Bocas del Toro, on October 18, 1963, was recovered at Algarrobo, 
Magdalena, Colombia, in February 1964 (Loftin, Rogers, and Hicks, 
Bird-Banding, 1966, p. 43). Although less common on the Caribbean 
slope, Wedel took several in San Blas at Permé and Obaldia between 
September 11 and May 10 (Griscom, Bull. Mus. Comp. Zool., vol. 72, 
19352, p.id7Z).. 

As a migrant, the Dickcissel is sometimes extraordinarily numerous. 
I have seen flocks containing thousands of birds. When flushed from 
the ground they resemble shorebirds, until they land in a tree; after 
ascertaining that all is safe they drop down to the ground again. They 
call constantly, giving twitters and more raspy notes. Skutch (A Nat- 


330 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


uralist in Costa Rica, 1971, p. 103) has heard them singing in winter. 
During the winter their distribution is more erratic and they are usually 
found in smaller groups, in clearings and marshes where they feed on 
seeds. Country people often call these birds “‘veinticuatro” from their 
habit of ranging in little flocks that popularly are believed always to 
number 24 individuals. A Dickcissel killed in an October rainstorm at 
Almirante weighed 24.36 g and had enough fat to travel another 846 
km (Rogers, Bird-Banding, 1965, p. 116). 


PSAROCOLIUS MONTEZUMA (Lesson): Montezuma Oropendola, 
Chacarero de Montezuma 


FIGuRE 27 


Cacicus Montezuma Lesson, 1830, Cent. Zool., livr. 2, p. 33, pl. 7. (Mexico.) 


Very large; narrow crest, head, upper back, throat, and breast black; 
wings and rest of body chestnut; central pair of tail feathers dark 
brown, others bright yellow; particolored bill. 

Description.—Length, male 395-475 mm, female 347-398 mm. Adult 
male, head, nape, upper back, throat, and breast black; rest of upper 
surface, including wing coverts, chestnut; primaries blackish, with 
basal half of outer web of eighth primary chestnut, and chestnut in- 
creasingly extensive on outer web of inner primaries; secondaries 
chestnut on outer web, blackish on inner web, increasingly reduced to- 
ward inner secondaries; central rectrices blackish, shorter than others, 
which are bright yellow; sides, flanks, and abdomen chestnut; central 
undersurface black with feathers tipped chestnut; underwing coverts 
black. The basal half of the bill is black, the terminal half is orange; a 
bare frontal shield is blackish; patch of bare skin below eye bluish and 
pink. 

Adult female, like male, but smaller and black more brownish on 
breast and rest of undersurface. 

Immature, like adult, but brown areas are duller; bill dusky at base, 
less yellow at tip. 

Measurements.—Males (10 from Panama, Costa Rica, Guatemala, 
Honduras, and Nicaragua), wing 218.8-261.0 (236.7), tail 170.9-200.2 
(181.6), culmen from base 71.5-81.1 (74.4, average of 9), tarsus 52.4- 
56.9 (54.5) mm. 

Females (10 from Panama, Costa Rica, Guatemala, Honduras, and 
Nicaragua), wing 187.3-208.2 (197.0), tail 146.0-163.5 (153.6), cul- 
men from base 57.4-61.1 (58.8), tarsus 43.8-46.9 (45.2) mm. 


FAMILY ICTERIDAE Sa 


Resident. Locally common in lowlands of the Caribbean slope in 
Bocas del Toro, less common farther east to the Canal Zone. It occurs 
as far north as eastern Mexico. In Bocas del Toro it has been recorded 
ptvoixaola (Peters, Bull: Mus. Comp. Zool., vol. 71, 1931, p. 343), 
Zegla, on the Teribe River (specimen collected by Austin Smith on 
May 16, 1927, now in the Havemeyer Collection at the Peabody Mu- 
seum, New Haven, Connecticut), Almirante, and at Changuinola and 
Isla de Colon, in Almirante Bay. Eisenmann states that “it seems to 
reach its regular southern limit in this area, although there are a few 
records from the Caribbean coast of the Canal Zone” (Condor, 1957, 
p. 257). In March 1952, I saw several at El Uracillo, Coclé, and Chilar, 
Colon. The Montezuma Oropendola was collected in the Canal Zone 
region by McLeannan (Salvin and Godman, Biol. Centr.-Amer., Aves, 
vol. 1 (pt. 52), 1886, p. 438) during the 19th century, but was not 
recorded there again until 1964 when N. G. Smith (in litt.) reported 
a colony in the Escobal Road area; in January 1969 Smith found a 
colony of 2 males and 12 females nesting in a big tree in a cow pasture 
on the Achiote Road southeast of the town of Achiote. This species 
can now be seen regularly on the Achiote and Escobal Roads and it has 
wandered to the Pacific side in the Rodman area of the southwestern 
section of the Canal Zone (Ridgely, 1976, p. 306). A specimen taken 
by M. J. Kelly in March 1921 at the Gaspar Savanna, near Chepo, 
eastern Province of Panama is now in the Everhart Museum, Scranton, 
Pennsylvania. Smith also reports (i litt. to Eisenmann) seeing a flock 
on April 9, 1975, above Santa Fé, Veraguas, at about 700 m on the 
Pacific slope and also over the Continental Divide. 


Fricure 27—Montezuma Oropendola, Chacarero de Montezuma, Psarocolius 
montesuma. 


338 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


The large Montezuma Oropendola feeds almost entirely on fruit and 
nectar, for which it travels considerable distances from its colonial 
nesting sites. Its flight 1s crowlike and direct. When feeding high in 
the tree tops this species is rather shy, and some of its food habits may 
have been overlooked; in Costa Rica, Larry Wolf (Wilson Bull., 1971, 
pp. 197-198) saw an oropendola catch and kill a fledgling Black-faced 
Grosbeak (Caryothraustes canadensis) that was part of a flock feeding 
in a tree. A male collected by Strauch (Bull. Brit. Orn: Claby 19779 p: 
64) weighed 528.5 g; a female weighed 241.8 g. 

This is a very vocal species. The male sings through most of the year, 
delivering a series of loud gurgling notes as he bows forward and raises 
his wings and then brings himself upright again. Once at El Uracillo 
I saw a male display by calling and opening his wings before his mate, 
who was on the branch beside him. He then moved over and touched 
her with his bill, finally pecking her on the rump; this was repeated 
several times. Both sexes also have several loud clucking and cacking 
calls. 

The Montezuma Oropendola nests in large and conspicuous colonies 
in Open areas in trees that are inaccessible to most predators. I have 
found active colonies during February and March in Bocas del Toro 
and western Colon. One colony on the Changuinola Canal in Almirante 
had approximately 25 nests on February 20, 1958. Females always 
considerably outnumber males. The Canal Zone area colonies are 
active from January to at least May, or sometimes early June; by July, 
nesting trees are deserted, and birds wander about in noisy bands. 

Skutch (Pac. Coast Avif., no. 31, 1954, pp. 287-304) has observed 
the nesting of this species in Costa Rica, where the season begins in 
January. One colony in a group of trees included 61 nests. The nests 
are placed on the outer twigs of trees from 13 to at least 33 m off the 
ground; they are hanging pouches 60 to 120 cm long and 18 to 23 cm in 
diameter near the bottom. It takes the female approximately 10 days, 
working alone, to weave the nest basket, which is made of long fibers 
of banana leaves, vines, and palm leaves, and she spends another 3 to 6 
days adding dead or dying leaves to the lining. Much of the nest ma- 
terial is brought from far away, but oropendolas also steal whatever 
they can from unattended nests. When not actually courting, the sexes 
ignore one another in the nest tree; the males provide no assistance in 
building or later in incubation or feeding the young, but they serve as 
sentries and accompany females gathering nest material. 

An egg in the British Museum collected in Costa Rica by C. F. Un- 
derwood was long and oval, dull buffy white with numerous scattered 


FAMILY ICTERIDAE 339 


scrawls and other irregular markings partly overlaid by shell so that 
they vary from dull chocolate and dull cinnamon to faint purple; it 
measured 43.8X26.3 mm. Three eggs measured by Skutch averaged 
38.1 24.3 mm. 

These oropendolas are sometimes parasitized by the Giant Cowbird 
(Scaphidura oryzivora). Although they attempt to drive the intruders 
out of their nest tree, a few cowbirds inevitably succeed in depositing 
an egg. Only some of the female oropendolas recognize the egg as 
foreign and remove it. 

The incubation period is 17 or 18 days (Smith, Nature, vol. 219, 
1968, p. 692) and the young remain in the nest approximately 30 days, 
during which time they are fed fruit. When they emerge, the young 
are quite large and able to fly a few hundred yards, even though they 
have never before had space to stretch their wings. The young are 1m- 
mediately led from the conspicuous and isolated nest tree to the safety 
of the forest. 

This species and P. guatimozinus have often been separated in the 
genus Gymnostinops. 


PSAROCOLIUS GUATIMOZINUS (Bonaparte): Black Oropendola, 
Chacarero Negro 


Ostinops guatimozinus Bonaparte, 1853, Compt. Rend. Acad. Sci. Paris, 37, p. 833. 
(Garrapata, middle Rio Magdalena, near Malena, Antioquia, Colombia.) 


Very large; few feathers of central crown elongated, forming a thin 
crest; all black except for dark chestnut scapulars, lower back, upper 
tail coverts, and abdomen, and bright yellow tail; particolored bill. 

Description.—Length, male 464-481 mm, female 387-396 mm. Adult 
(sexes alike except for size), entire body black except middle of back 
to upper tail coverts, abdomen, and undertail coverts dark chestnut, 
darkest on rump and upper tail coverts; wing coverts dark chestnut, 
remiges black; central pair of rectrices black, shorter than others, 
which are bright yellow; underwing coverts black. 

A male collected January 31, 1964, at Pucro, Darién had the iris 
light wood brown; swollen cheek patch deep clear blue; bare skin be- 
hind eye paler blue; lower edge of lores like cheek patch; bare forehead 
reddish orange, changing to orange from most of center; top of bill 
orange-red; rest of bill black; top of tongue pale gray; inside of mouth 
black except for tip, this orange-red, shading distally to yellowish on 
mandible; bare skin on side of throat dull red, bordered narrowly by 
bluish white, this color extending on either side of central feathering 
inside rami; tarsus and claws black; underside of toe pads dull yellow, 


340 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


blacker on distal pads. Two females taken at Pucro on February 2, 
1964, had the soft parts like the male’s except that the bare forehead 
was dull red. © 

Measurements.—Males (7 from Darién and Colombia), wing 241.0- 
264.0 (253.9), tail 184.5-201.0 (195.3), culmen from base 65.2-71.2 
(68.1, average of 6), tarsus 60.0-63.6 (55.8) mm. 

Females (9 from Darién and Colombia), wing 198.0-228.0 (205.8), 
tail 154.6-180.0 (168.5), culmen from base 51.6-67.7 (57.1, average of 
8), tarsus 43.7-59.3 (48.6) mm. 

Resident. Fairly common in the lowlands of Darien, and may range 
west into the upper Bayano River valley in eastern Province of Pan- 
ama, where there are some recent unconfirmed sightings (Ridgely, 
1976, p. 306) that may be referable to Crested Oropendolas. The Black 
Oropendola is also found in northern Colombia. In Darién it has been 
collected at Marraganti by E. A. Goldman in 1912, at Tapalisa, Chepi- 
gana, and E] Real on the Rio Tuira by W. B. Richardson in 1914 and 
1915, and on the Rio Sambu by the Fifth George Vanderbilt Expe- 
dition in 1941. Other Darién specimen records (fide Eisenmann) are 
from Santa Fé, Rio Chucunaque, and Rio Pirre. 

In 1964 I found a group of males at a nest colony at Pucro, Darién, 
in late January. The colony was in a tree at the border of a stand of 
platanos. Near or at the nest, the males called hollowly and then fell 
forward to hang upside down briefly. This was varied by the hollow 
calls being followed by high pitched excited notes—kwee, kwee, kwee 
or keea, kee-a, kee-a, often uttered as the bird dashed away in sudden 
flight. The colony’s 20 or so nests were mostly complete and were 
grouped closely in the ends of two or three branches. On the ground 
were the nests of last season that apparently had been torn down by the 
birds when they started the new construction; many of them were com- 
plete. One that I examined had many small, partly cut up dried leaves 
in the bottom. 

A Black Oropendola that I watched feeding in a flowering tree at 
Pucro thrust its bill into a closed flower and gaped to open the corolla. 
I was interested to find when skinning one of these birds that the bare 
blue patches, projecting roundly, and hard to the touch, covered a jaw 
muscle of similar size. 

Females that I collected in early February were nearly in laying con- 
dition, but in the evenings all the birds I saw were flying to roosts 
rather than remaining at the nest tree. 

Two eggs from Remedios, Antioquia, Colombia, in the Salvin- 
Godman collection at the British Museum are oval, dull white, with 


FAMILY ICTERIDAE 341 


the shell very slightly roughened, have a faint gloss, and are marked 
with a few widely scattered irregular spots that vary from cinnamon 
and chocolate to purple, scattered among which are many tiny dots of 
the same color. They measure 34.0 X 24.8 and 35.2 x 24.8 mm. 


PSAROCOLIUS WAGLERI (Gray): Chestnut-headed Oropendola, 
Chacarero Comun 


FIGURE 28 


C. [acicus] Wagleri G.R. Gray, 1845, Genera, Birds, 2, p. 342, pl. 85. (“Coban,” 
Guatemala. ) 

Zaryhynchus wagleri ridgwayt van Rossem, 1934, Bull. Mus. Comp. Zool., 77, p. 
405. (Limon, Limon, Costa Rica.) 


Large; few narrow feathers of central crown elongated, forming a 
thin crest; head, rump, flanks, and abdomen dark brown; wings and 
rest of body black; tail bright yellow. 

Description.—Length, male 336-353 mm, female 273-289 mm. Adult 
male, head, nape, throat, and upper breast bay; rump, upper tail coverts, 
flanks, lower abdomen, and undertail coverts chestnut; rest of body and 
wings iridescent black; central pair of rectrices (slightly shorter than 
others) and outer web of outermost pair black, others bright yellow; 
underwing coverts black. 

Adult female, like male, but smaller, and less glossy black (browner) 
below. | 

A male collected at El Copé, Coclé, on February 23, 1962, had the 
iris pale blue, tip of bill extending back on cutting edge of mandible 
and on base of rami and around base of pileum light neutral gray; rest 
of bill yellowish buff; tarsus, toes, and claws black. Some males have 
the bill greenish yellow with a dusky tip (Eisenmann). A female taken 
at Puerto Armuelles, Chiriqui on January 30, 1966, had the iris pale 
blue; bill light greenish yellow changing to pale neutral gray at tip; in- 
side of mouth including top of tongue pale neutral gray; tarsus and toes 
fuscous-brown; claws black. 

Measurements.—Males (10 from Panama), wing 187.0-227.0 
(214.1), tail 124.8-144.0 (133.5), culmen from base 62.4-72.4 (66.1), 
tarsus 37.7-41.1 (40.0) mm. 

Females (10 from Panama), wing 145.4-161.9 (152.1), tail 100.0- 
117.3 (110.0), culmen from base 46.6-52.0 (49.9), tarsus 30.1-33.8 
(31.8) mm. 

Resident. Common but local, in the humid lowlands of both slopes 
in forests, second growth, borders, and open areas that have trees suit- 
able for nesting. More local on the Pacific slope, where it is absent en- 


342 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


tirely from the drier parts of the Azuero Peninsula and from southern 
Coclé. This species has been recorded in the foothills as high as 1350 
m (Ridgely, 1976, p. 306); Frank Hartman collected a female at El 
Volcan (1230 m), Chiriqui (now in the collection of Ohio State Uni- 
versity ) and I have seen them feeding in a banana plantation at 1200 m 
at Santa Clara, Chiriqui. 


Figure 28.—Chesnut-headed Oropendola, Chacarero Comin, Psarocolius waglen, 
male. 


The populations from Nicaragua southward have in the past been 
recognized under the name ridgwayi and are supposed to be charac- 
terized by paler coloration. The series in the Smithsonian collection 
indicates that if there is a point of demarcation between northern and 
southern populations, it is somewhere south of Panama, whereas birds 
from Mexico through Panama do not show any geographic variation. 
Therefore, ridgway1 becomes a synonym of wagleri. There is as yet 
insufficient material from South America for further nomenclatural 
action. 

This is the most widespread oropendola in Panama. Its nest colonies, 
sometimes with as many as a hundred long, globular pouches hanging 
from an isolated tree or grove, often on the side of a well-traveled 
highway or even in front of a building (as at Coco Solo Hospital), are 


FAMILY ICTERIDAE 343 


a familiar sight. Like other oropendolas, this is a highly vocal bird. 
Eisenmann (Smiths. Misc. Coll., vol. 117, no. 5, 1952, p. 53) lists some 
of its calls as a “liquid croak, grwackoo, grwackoo, grwak, grwak; 
often grwackoo alone; also a series of gurgling notes like the sound of 
dripping water, plup, plup, plup, plup-looupoo; also plup alone.” Eisen- 
mann also reports watching a male giving a liquid song, 00-lo0-ooh, tloo, 
coo-00, tloo-coo-oo, in late April. Chapman (Bull. Amer. Mus. Nat. 
Hist., vol. 48, art. 3, 1928, pp. 132-133) lists several other calls. Males 
also produce a variety of noises with their wings in flight that are part 
of courtship displays. 

The Chestnut-headed Oropendola feeds high in trees. It often moves 
in groups to and from the nesting colony, and during the nonbreeding 
season moves through the forest in noisy flocks. One collected by E. A. 
Goldman at Cana, Darién, on June 19, 1912, contained acridid frag- 
ments 38%, elytra of an otiorhynchid 2%, about 550 seeds and 2 other 
indeterminate 60%. The large mass of seeds was apparently from 
berries. Leck (Auk, 1974, p. 163) has observed this species feeding on 
the nectar of the large white flowers of the balsa tree (Ochroma limon- 
ensis). Two males collected by Strauch (Bull. Brit. Orn. Club, 1977, 
p. 64) weighed 213.4 and 239.4 g; 2 females weighed 134.2 and 141.3 ¢g. 

At Barro Colorado Island, Canal Zone, where Chapman (op. cit. pp. 
123-166) investigated the nesting habits of this species in great detail, 
birds return to a previously used nest tree in late December for brief 
inspections, and females begin constructing new nests in early Janu- 
ary. In 1964, at Tacarcuna Village in Darién I founda colony on March 
8 to which pairs came occasionally to inspect and peer into old nests, 
usually visiting several in turn. By March 12 they came in groups of 
12 or 15 and there was activity in the nest tree all day, although still 
no new construction. At Barro Colorado most breeding activity ended 
by the beginning of the rainy season in May, although a few nests were 
active into July. The nest tree is always chosen for its situation beyond 
the reach of most predators. In Darién I found a group of nests swing- 
ing in a tree over the Rio Imamado and another at Jaqué on a tree 
whose trunk had a large wasps’ nest. Usually a colony contains 25 to 
50 active oropendola nests. The nests rarely survive from year to year 
in usable condition, so new ones are built each season. They are placed 
on the outermost twigs that are about the diameter of a pencil in width; 
most nests are on the leeward side of the tree, to avoid damage from 
the trade winds that blow during this season. The nests seem to be 
placed as near to one another as possible. 

Male oropendolas go through a noisy and aggressive courtship peri- 


344 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


od, during which several may pursue a single female until one has estab- 
lished his dominance and the other males look elsewhere. At the Barro 
Colorado colony Chapman studied, females outnumbered males about 
6to 1. Males may accompany particular females as they go off to gather 
nesting material, but as soon as incubation begins the males loose in- 
terest. 

The nest itself is a long pouch woven out of lengthy rootlets, tendrils, 
and fibers that are gathered in the forest or stolen from neighbors. 
Chapman found nests that measured from 55 to 100 cm in length; the 
diameter of the pouch was always 19 cm. At the beginning of the sea- 
son most nests take about 31 days to complete; later, however, they are 
built more rapidly. The two eggs that form a clutch are “pale blue with 
numerous irregular shaped brownish-black marks varying in size from 
a pinpoint to a currant and clustering most thickly about the larger 
end” (Chapman, op. cit., p. 147). The one egg Chapman was able to 
measure was 33 by 22 mm. 

The incubation period is 17 days and the young emerge from the nest 
approximately 33 days after hatching. Incubation and care of the young 
are performed entirely by the female. The oropendolas have two avian 
parasites to contend with, the Giant Cowbird (Scaphidura oryzwwora) 
and the Piratic Flycatcher (Legatus leucophaius). The cowbird is rec- 
ognized as an enemy from the time the female cowbirds first visit the 
colony tree, when nests are still under construction; groups of oropen- 
dolas attempt to drive them away, but inevitably a few cowbird eggs 
are deposited in oropendola nests. Chapman mentioned no incidents of 
oropendolas recognizing the strange egg and removing it. The Piratic 
Flycatcher attempts to harass at least 1 female oropendola to the point 
where she deserts her nest, which the pair of flycatchers then take over 
and use as their own. In 1928 a pair of these flycatchers so distracted 
the birds Chapman was observing that the entire colony deserted its 
tree: 

This species has often been separated in the genus Zarhynchus. 


PSAROCOLIUS DECUMANUS MELANTERUS (Todd): Crested 
Oropendola, Charcarero Crestado 


Ostinops decumanus melanterus Todd, 1917, Proc. Biol. Soc. Washington, 30, p. 3. 

(Las Vegas, Santa Marta, Colombia.) 

Large; a few feathers of central crown elongated, forming a slight 
crest; all black except for chestnut rump and undertail coverts, and yel- 
low tail. Bill, extended to form a frontal shield, pale yellowish green 
or waxy yellow, without dusky tip. 


FAMILY ICTERIDAE 345 


Description.—Length, male 412-442 mm, female 319-340 mm. Adult 
(sexes alike except for size), entirely black except for rump, lower 
abdomen, and upper and undertail coverts, which are chestnut, and for 
tail, in which all but central pair of rectrices are bright yellow; central 
pair of rectrices black, shorter than others. 

Immature, similar, but black and chestnut areas duller, sometimes 
with brown edging; bill yellowish brown or brownish white. 

A female collected at El Real, Darién, on January 23, 1964, had the 
iris dark gray; bill ivory, slightly more yellowish on basal third; tarsus, 
toes, and claws black. A male taken at Alanje, Chiriqui, on March 12, 
1960, had the iris bright blue. 

Measurements.—Males (10 from Panama and Colombia), wing 
196.0-236.0 (213.7), tail 170.2-207.0 (188.0), culmen from base 57.9- 
64.8 (60.4), tarsus 49.3-51.4 (50.6) mm. 

Females (10 from Panama), wing 160.0-179.8 (168.6), tail 137.1- 
159.0 (151.6), culmen from base 46.0-49.3 (47.7), tarsus 39.0-43.3 
(41.3) mm. 

Resident. Locally common in forest borders, second-growth wood- 
lands, and clearings in the lowlands of the Pacific slope from western 
Chiriqui (Buguba, Divala, Concepcion), where it is scarce with few 
recent reports, to the Colombian boundary in Darien, including the 
Azuero Peninsula (south to Tonosi and Pedasi, Los Santos), ranging 
at least locally to the lower edge of the Subtropical Zone (above El 
Valle, Coclé; Cana, Darién). In March 1981 it was the only oropendola 
present at Cana; here it was abundant and nesting (Ridgely, in litt.). 
On the Caribbean side it is found in scattered colonies along the Rio 
Indio, from above the mouth near Chilar, Colon to El Uracillo, Coclé, 
and in the Canal Zone and vicinity (Ridgely, 1976, p. 307). This 
species is much commoner in Darién than it is further west, and al- 
though unrecorded from San Blas, will probably be found there as it is 
common in adjacent northern Colombia. In the Bayano Valley of 
eastern Province of Panama, however, it is still outnumbered by P. 
waglert. Other races of Psarocolius decumanus are found in Colombia 
east of the Andes, south through Ecuador, Peru, and Bolivia to north- 
ern Argentina and east to the Guianas, Trinidad and Tobago, and much 
of Brazil. 

The habits of this species are very like those of other oropendolas. 
Its calls are similar in quality to the vocalizations of P. wagleri, al- 
though they seem less varied and most are lower in pitch. It nests in 
colonies, often in smallish Cecro pia trees, sometimes in association with 
P. wagleri or with Yellow-rumped Caciques (Cacicus cela). A colony 


346 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


I found at Chilar, Colon, on March 7, 1952, had 15 or 20 nests in a tree 
over the Rio Indio. Their baglike nests give the oropendolas the local 
name of chacarero, because the nest suggests the chacara, a knit bag 
used by the country people. Salvadori and Festa (Bol. Mus. Zool. 
Anat. Comp. R. Univ. Torino, vol. 14, no. 339, 1899, p. 5) noted that a 
native name in Darién was uropendulo. 

The nesting season in Panama seems to begin later than that of other 
oropendolas. On March 9, 1951, I shot 2 females at La Campana, Pan- 
ama, that had ovaries beginning to develop. On March 22, 1949, I saw 
females looking at nesting sites at Chico, Panama. I saw females carry- 
ing food to young in the nest on April 22, 1949, at Chepo, Panama. 
In March 1981, Ridgely (2m litt.) found nests under construction in 
Cana, Darién. In the Canal Zone the nesting season extends into late 
June (Ridgely, im litt.). Courtship and nest construction by the race 
P. d. insularis on Trinidad are discussed in great detail by Drury (Zoo- 
logica, 47, pt. 1, 1962, pp. 39-58). A photograph in Nature (vol. 219, 
1968, p. 690) shows a light-colored egg with a few dark marks. Schafer 
(Bonner. Zool. Beitr., 1951, pp. 1-148) writes that in Venezuela eggs 
were white, pale gray, or pink with dots and spots of black irregularly 
scattered over the larger end. Two eggs measured 23X34 mm and 
each weighed 12 g. The clutch was 1 or 2. 

When preparing specimens of this oropendola I noted that the oil 
gland was very large and contained considerable thickened fluid which 
I did not encounter in any other oropendola. Two females collected by 
Strauch (Bull. Brit. Orn. Club, 1977, p. 64) weighed 178.3 and 191.8 g. 

In July 1975, Ridgely (im litt.) found a roost at Matuganti (above 
the mouth of the Rio Pucro) that contained 2000-2500 oropendolas 
and caciques. Of these, probably over 1000 were Crested Oropendolas, 
with Yellow-rumped Caciques, Black and Chestnut-headed Oropen- 
dolas, and Giant Cowbirds in diminishing proportions. 


CACICUS CELA VITELLINUS (Lawrence): Yellow-rumped Cacique, 
Charro de Rabadilla Amarilla 


Cacicus vitellinus Lawrence, 1864, Proc. Acad. Nat. Sci. Philadelphia, p. 107. 
(“New Greneda, Isthmus of Panama.” ) 


Rather large; all black except for bright yellow lower back, rump, 
undertail coverts, and small patch on wing coverts; bill pale yellow. 

Description.—Length, male 268-292 mm, female 223-233 mm. Adult 
male, body all black except for lower back to upper tail coverts, lower 
abdomen, and undertail coverts, which are rich yellow to orange-yellow; 


FAMILY ICTERIDAE 347 


wings black with patch of rich yellow on central coverts, often partially 
concealed; tail black with basal quarter of feathers paler yellow. 

Adult female, like male, but smaller, and black duller, tinged with 
olive below. 

A male collected at Cafita, Province of Panama, on February 7, 
1962, had the iris light blue; bill dark ivory-yellow with a very narrow 
line of neutral gray around entire base, faintest on lower border of 
mandibular rami; tarsus, toes, and claws fuscous-black. A female 
taken at Juan Mina, Canal Zone, on January 11, 1961, had the iris light 
blue; bill yellow, shading to very pale grayish white at tip; tarsus and 
toes fuscous-brown. 

Measurements——Males (10 from Panama), wing 168.5-183.0 
(lore), tail) 10915-118)5\ (115.2), culnmien from base 36,0-40:5 (39.1), 
tarsus 31.4-34.3 (33.2) mm. 

Females (10 from Panama), wing 120.0-140.1 (130.8), tail 88.5- 
97.0 (92.9), culmen from base 31.6-34.2 (32.5), tarsus 27.5-32.2 (29.4) 
mm. 

Resident. Fairly common in forest, second-growth woodlands, 
borders, and clearing in the lowlands. Most numerous in Darién and 
the Rio Bayano Valley of eastern Province of Panama. On the Pacific 
slope found from Sona (Bangs, Proc. New England Zool. Club, vol. 3, 
1902, p. 64) and Paracote (Aldrich, Scient. Publ. Cleveland Mus. Nat. 
iniseevoles, 1937,). 127), Veraguas, east to Colombia, and on the 
Caribbean slope eastward from the Rio Indio in western Colon. In the 
lowlands of the Canal Zone area, it extends across the Isthmus, and in 
Darién it has been collected as high as 540 m by FE. A. Goldman at 
Cana, on May 22, 1912. In March 1981, Ridgely (in litt.) found it at 
Cana up to 700 m. This race is also found in northern Colombia; other 
races occur as far south as northern Bolivia and central Brazil. 

The Panama form, vitellinus (Lawrence’s Cacique), was formerly 
considered specifically distinct from the Amazonian and Guianan C, 
cela cela, from which it differs in the lesser extent of, but richer, yellow 
in the wings and tail. The nominate race is well known throughout its 
range as a mimic of other birds and of the sounds of mechanical ob- 
jects; this is unreported for vitellinus. In Venezuela nominate cela 
is often caged because of its vocal abilities. 

The Yellow-rumped Cacique resembles the oropendolas in many 
ways, and sometimes nests in the same trees with them, although usually 
in a group somewhat removed. Like the oropendolas, it feeds high in 
trees, where it is often difficult to observe. Once at Cafita, Panama, I 
was able to see a flock well enough to note that the birds were eating 


348 BIRDS OF THE REPUBLIC OF PANAMA—PART A 


figs. IX. A. Goldman recorded the stomach contents of 2 that he col- 
lected: one contained 11 caterpillars, 9 of one species, 2 of another 55%, 
25 seeds with bits of red fruit pulp surrounding them 45%; the other 
had bits of 2 ants 10%, a large otiorhynchid near Lachnopus 10%, re- 
mains of two other weevils 5%, elytron of another beetle trace, 58 
seeds undetermined, a few other smaller ones, and a mass of vegetable 
fiber 75%. A male collected by Strauch (Bull. Brit. Orn. Club, 1977, 
p. 64) weighed 94.9 g. 

Caciques call constantly with a series of highly varied conversational 
notes; these are closely similar to those of Psarocolius wagleri, but with 
fewer bubbling calls and explosive notes, and, in all, more musical. 
Some that I noted are wick-a-weo, char-che-ar, chock-chu-chou, wicka- 
wicko, and chut-chu-chut. The female seems limited to liquid chirps 
and clucks. 

Although they sometimes nest with oropendolas, the cacique’s breed- 
ing season appears to start somewhat later. It may depend on locality 
and the rainfall or dryness of a particular year. On March 9, 1957, at 
Pedasi, Los Santos, I watched a male displaying before a female 
perched beside him; the male arched his neck and raised the feathers 
there and on the rump and gave his gabbling calls. During April of 
1949, I found several colonies in the Province of Panama that were 
still under construction: near Cerro Azul I saw a group starting a new 
colony in a tree in recently cleared fields at San Miguel on April 19, 
and at Chepo I watched females carrying nest material to a colony on 
April 26. Skutch, who studied this species in 1935 at Barro Colorado 
Island in the Canal Zone (Pac. Coast. Avif. no. 31, 1954, pp. 305-315) 
found them building nests there during April and early May. 

The nest is a hanging pouch woven from strips of palm fronds and 
fibrous strands of leaves and vines, and is lined with softer fibers. It 
is 30 to 45 cm long, generally shorter and more oblong than oropendola 
nests. Eisenmann points out that the nests are placed closer together, 
often touching each other. In South America caciques have evolved an 
interesting adaptation that makes nesting in the rainy season less dif- 
ficult: when the rains begin in the middle of the breeding cycle the fe- 
male weaves a roof over the nest entrance so that the interior remains 
dry (Cherrie, Mus. Brooklyn Inst. Arts and Sci. Bull., 2, 1916, p. 204). 
This has not been observed in Panama, but it is certain that Yellow- 
rumped Caciques have not finished raising their young by the beginning 
of the rainy season. When not in a tree surrounded by a clearing, the 
colony is sometimes over water, as the 1 Skutch watched and 1 that I 
found at Chepo that consisted of 15 nests hung from a pendant branch 


FAMILY ICTERIDAE 349 


2 or 3 m above the Rio Mamoni. Very frequently a tree containing a 
wasp colony is chosen; Cecropias that host stinging ants are sometimes 
used. At the Barro Colorado colony there were 8 males and 15 females; 
as in the oropendolas, the females build the nest and raise the young 
independently. 

The eggs of C. c. vitellinus have not been described, but those of the 
nominate race in Venezuela are “white with a faint bluish wash, marked 
with specks, spots and blotches of chestnut over vinaceous brown” 
(Cherrie, op. cit., p. 205); in Venezuela the clutch consists of two eggs. 
The incubation period is 16 days (Smith, Nature, vol. 219, 1968, p. 
692). The first eggs at Skutch’s colony hatched on May 12; 3 days later 
over half the nests had young. The colony, however, was pillaged by a 
snake (Spilotes pullatus), which destroyed the contents of all but 2 
nests. Less severe nuisances endured by the colony were the parasitic 
Giant Cowbird (Scaphidura oryzivora), although Skutch never saw 
one actually entering a nest, and the Piratic Flycatcher (Legatus leu- 
cophaius ), which harassed the caciques but never took over one of their 
nests. 


CACICUS UROPYGIALIS (Lafresnaye): Scarlet-rumped Cacique, 
Charro de Rabadilla Escarlata 


Cassicus uropygialis Lafresnaye, 1843, Rev. Zool. [Paris], 6, p. 290. (“Bogota,” 
Colombia.) 


Medium size; all black with scarlet rump. 

Description.—Length 200-235 mm. Adult (sexes alike), all black, 
with scarlet rump and lower back. 

Juvenile, like adult, but red rump duller, brownish orange. 

These are forest birds, found in pairs or groups of half a dozen or 
more, that range mainly through the higher branches. Skutch, who has 
observed them in Costa Rica (Pub. Nutt. Orn. Club no. 10, 1972, pp. 
173-181) noted that there they were often loosely associated with other 
treetop birds, including oropendolas, Black-faced Grosbeaks (Caryo- 
thraustes poliogaster), Purple-throated Fruitcrows (Querula pur pur- 
ata), and White-fronted Nunbirds (Monasa morphoeus). Much of 
their movement is behind leaf cover, so that often it is not easy to dis- 
tinguish them from related species of caciques. The light-colored bill 
is usually prominent, but aside from this they appear plain black, as the 
red of the lower back is ordinarily covered by the wings. They often 
whistle musically as they move about, and display considerable curio- 
sity. At any unusual circumstance they may come with flitting wings 
and jerking tails to call and scold vociferously near at hand. They feed 


350 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


on insects, fruit, and nectar. Four males collected by Strauch (Bull. 
Brit. Orn. Club, 1977, p. 64) weighed from 62.5 to 71.7 g; 5 females 
weighed from 50.0 to 57.6 g. 

Although while they range in small groups or pairs during the day, 
at night they tend to gather to sleep in roosts. On the slopes of Cerro 
Pirre in Darién in late January and early February of 1961, several 
hundred came to tall trees bordering an open marsh near our camp at 
450 m elevation on the upper reaches of the Rio Seteganti. Small bands 
began to arrive after sunset to perch in the high tree crown where they 
moved about restlessly until finally settled for the night. In the morn- 
ing they began to call as soon as it was light, and then flew about with 
noisy wings through the fog-screened treetops until the sun appeared, 
when they spread out into the adjacent forest. Another roost in early 
March was located in a band of tall trees along the mouth of the Que- 
brada Candelaria, in the Province of Panama, where that stream joins 
the Rio Pequeni. Here the birds assembled in the forest behind the 
clearing and came to the roosting area in groups of 25 to 30. These 
swept in over the tops with much calling and wing noise, rising and 
falling in undulating lines. 

This species ranges from Nicaragua south through Costa Rica, Pan- 
ama, Colombia, and Ecuador to northern Peru. Three subspecies are 
recognized currently, of which 2 (microrhynchus and pacificus) are 
found in Panama. These differ from the typical race, Cacicus uropy- 
gialis uropygialis, by their smaller size. The race microrhynchus has 
often been treated as a separated species, C. microrhynchus, but paci- 
ficus seems intermediate in bill shape and geographic position. 


CACICUS UROPYGIALIS MICRORHYNCHUS (Sclater and Salvin) 


Cassiculus microrhynchus Sclater and Salvin, Proc. Zool. Soc. London, 1864 (Feb., 
1865), p. 353 (Lion Hill, Canal Zone.) 


Characters._Compared with C. u. pacificus, bill, particularly the 
culmen, more slender, with the mandibular rami normal in form, not 
swollen laterally; concealed white bases of feathers of dorsal surface 
found at base of hindneck only, in some specimens absent. 

A male collected at Juan Mina, Canal Zone, on January 14, 1961, 
had the iris pale blue; bill light yellow at base changing to greenish near 
center and continuing thus to tip; tarsus and toes fuscous-black. Fe- 
males | have collected were similar. 

Measurements.—Males (10 from Panama), wing 122.0-136.5 


FAMILY ICTERIDAE 351 


(129.0), tail 83.3-96.5 (89.7), culmen from base 28.8-30.2 (29.6), tar- 
sus 27.4-30.0 (28.8) mm. 

Females (10 from Panama), wing 109.7-117.3 (113.9), tail 77.5- 
O75) (00.4), culmen from base 25.7-29.2'(27.5)} tarsus 25.7-27.8 
(26:9) mm. 

Resident. found in forested areas in the lowlands of both slopes 
from Chiriqui and Bocas del Toro to the eastern section of the Province 
of Panama (Utivé) and eastern San Blas (Permé, Puerto Obaldia), 
ranging to 900 m in mountain areas (Santa I*é, Veraguas). To the 
west and north this race ranges through Costa Rica to Nicaragua. In 
Panama it is absent from the savanna areas of the Pacific slope and 
there are no records from the Azuero Peninsula. Felling of the original 
forest has greatly reduced their range in Chiriqui, where Brown secured 
specimens at Divala in November and December, 1900 (Bangs, Auk, 
1901, p. 370), and on the Pacific slope of Veraguas, where Arcé found 
it near Santiago (Salvin and Godman, Biol. Centr.-Amer., Aves, vol. 1 
(pt. 53), 1886, p. 442). There are specimens from Puerto Armuelles 
and “Baru,” Chiriqui, in the California Academy of Science taken by 
M. EK. Davidson in November 1929. There appear to be very few, per- 
haps none, left in these areas today, although the bird is still fairly 
common on the Caribbean slope. Ridgely (in litt.) is unaware of any 
reports from Chiriqui or Pacific Veraguas other than above Santa Fe, 
where several groups were seen in January 1974 at about 900 m. In 
April 1980, Ridgely was struck by their abundance in the forests of the 
Changuinola River in western Bocas del Toro. 

On March 25, 1949, I found a mated pair at Utivé, in the foothills 
Oiene @erro Azul, Panama, and Arbib and Loetscher (Auk, 11935, p. 
237) included this as one of the species noted as breeding in or near the 
Canal Zone in July and August. On March 25, 1975, E. S. Morton and 
IXisenmann saw a female on Achiote Road, Canal Zone, building a nest 
hung from a tree branch, while in the same tree a male perched about 
7maway. On June 25, 1952, Eisenmann and J. Bull found a nest be- 
ing built on a vine growing on a bridge between Fort Sherman and Fort 
San Lorenzo, Canal Zone. 

Skutch (Pub. Nutt. Orn. Club, no. 10, 1972, pp. 173-181) has studied 
the species in Costa Rica, where he found that, unlike Cacicus cela and 
the oropendolas, C. uropygialis is monogamous and nests solitarily; 
other characters it shares with Jcterus rather than the larger species are 
singing by the female and feeding of the nestlings by the male. N. G. 
Smith (in litt. to Eisenmann) found that birds about Almirante, Bocas 


352 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


del Toro, on the Rio Ciri Grande, Province of Panama, and apparently 
in western Panama east to the western part of the Canal Zone all nest 
solitarily, while in the eastern part of the Canal Zone (Madden Forest 
Preserve) eastward, colonial nesting apparently prevailed, although 
some solitary nesting occurred. 

The nest is a long pouch—one measured 64 cm in length—placed in 
a tree from 4 to 33 m from the ground; often a tree containing a wasp 
colony is selected. Skutch found birds building nests in late April and 
early May. One nest contained two eggs on April 24; they were “white 
marked with a few light brown and blackish spots and some short 
scrawls on the thicker end, still fewer elsewhere.” Incubation is per- 
formed exclusively by the female. At one nest two eggs hatched by May 
1; the young had pink skin, no down, and mouth interiors of pale pink. 
At 3 or 4 days their pinfeathers were sprouting and their eyes were 
partly open. The young were fed insects;- unfortunately this nest was 
destroyed, so that Skutch could not observe their growth. Young, well- 
grown birds still accompany their parents and are fed by both of them; 
Skutch found one such family in mid-March, when a juvenile was be- 
ing fed by both parents and had little success finding food on its own. 

After breeding, these solitary nesters often gather in bands. 

The voice of C. u. microrhynchus is far more musical than that of 
the Yellow-rumped Cacique, C. cela; it consists chiefly of loud, clear 
whistles. On June 23, 1952, Eisenmann noted a vocalization as a rather 
human-sounding whistle wheew-whee-whee-whee-wheet, sometimes ab- 
breviated to wheew-whee-wheet. Characteristic also was a rather sweet, 
tremulous, though burry, or throaty, whistle, ree-oo0, ree-oo varied to 
kreéoo or shreew-shreew. 

An occasional adult male of this race shows a faint swelling on the 
outer face of the base of the mandibular rami, an indication of approach 
to the condition found in C. u. pacificus, but this is not usual. The 2 
races are similar in size. 


CACICUS UROPYGIALIS PACIFICUS Chapman 


Cacicus uropygialis pacificus Chapman, Bull. Amer. Mus. Nat. Hist., vol. 34, Dec. 
30, 1915, p. 657. (Alto Bonito, Rio Sucio, Antioquia, Colombia.) 


Characters.—Bill larger, with the bases of the mandibular rami dis- 
tinctly swollen; concealed white on bases of the feathers of the dorsal 
area much more extensive, both in distribution and on the individual 
feathers. 

Measurements.—Males (10 from Darién and Colombia), wing 


FAMILY ICTERIDAE 353 


122.2-137.0 (132.2), tail 87.8-94.9 (91.1), culmen from base 28.2-31.0 
(29.6), tarsus 28.2-31.5 (29.5) mm. 

Females (8 from Darién and Colombia), wing 113.1-122.5 (118.6), 
tail 76.0-85.2 (81.0), culmen from base 26.6-28.9 (27.9), tarsus 24.5- 
Zoe (26.5) mm: 

Resident. Fairly common in eastern Darien, where it has been re- 
corded from the mouth of the Rio Pifias on Bahia Pinas, Jaqué, Rio 
Jaqué, Cerro Quia, and Cerro Pirre; found also in western Colombia 
and in western Ecuador. From the somewhat scanty data, there may 
_bea gap between the range of this form and that of C. u. microrhynchus. 
On the Rio Jaqué I once found a flock of a half dozen feeding low in 
dense growth along the river. A male that I collected at Jaqué on April 
13, 1946, was in breeding condition. 

The vocalizations of this race include a whistled teeo or keeo, with- 
out the burry quality of the corresponding call of microrhynchus in the 
Canal Zone (Eisenmann, in litt.). 


CACICUS HOLOSERICEUS HOLOSERICEUS (Deppe): Yellow-billed 
Cacique, Charro Piquihueso 


Sturnus holosericeus W. Deppe, 1830, Preis-Verz. Saug., Vogel, etc., Mexico, 
pl. 1. (Alvarado, Veracruz, Mexico.) 

Amblycercus holosericeus centralis Todd, 1916, Proc. Biol. Soc. Washington, 29, 
p. 95. (Rio Sicsola, Costa Rica.) 


Medium size; entirely black; bill yellowish. 

Description.—Length 202-222 mm. Adult (sexes alike), entirely 
black. 

A male collected at Fl Llano, Province of Panama, on February 4, 
1962, had the iris orange; bill light yellowish green, except the nasal 
operculum and extreme base of maxilla on the side immediately below, 
which are neutral gray; a narrow line of neutral gray along side of 
mandible also; tarsus, toes, and claws neutral gray. A female taken 
the same day was similar. 

Measurements.—Males (10 from Panama), wing 92.0-105.6 (98.8), 
tail 88.5-101.2 (94.9), culmen from base 28.7-31.7 (30.6), tarsus 29.8- 
33.8 (32.3) mm. 

Females (10 from Panama), wing 85.9-96.8 (90.7), tail 84.0-99.0 
(91.4), culmen from base 25.4-29.2 (27.3), tarsus 27.1-31.8 (30.3) 
mm. : 

Resident. Locally common in thickets of wooded borders, clearings, 
and second growth throughout the lowlands on both slopes of the main- 
land and more irregularly found in the foothills and lower highlands. 


354 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


Monniche (Blake, Fieldiana: Zool., vol. 36, no. 5, 1958, pp. 563-564) 
collected it on the Volcan de Chiriqui between 1590 and 2910 m and 
W. W. Brown, Jr., found it there between 2850 and 3090 m (Bangs, 
Proc. New England Zool. Club, vol. 3, 1902, p. 64). In Coclé it has 
been collected at 750 m at El Valle, and in Darién at 600 m at Cana. I 
have not found it in the lowland scrub of the drier eastern side of the 
Azuero Peninsula, though it occurs in the more humid western slope 
in this area, where Aldrich and Bole (Scient. Publ. Cleveland Mus. 
Nat. Hist., vol. 7, 1927, pp. 24-25) considered it common in semi- 
deciduous coastal forest near Montijo Bay, Veraguas, and uncommon 
in rain forest between 300 and 900 m. This race also occurs north to 
central Mexico and south to northern Colombia; other races are found 
south to eastern Peru and Bolivia. | 

The race centralis of Costa Rica and Panama, described by Todd 
(Proc. Biol. Soc. Wash., 1916, 29, p. 95) as having the wings longer 
than the tail, rather than the reverse, as supposedly found in popula- 
tions farther north, is not valid. Both Peters (Bull. Mus. Comp. Zool., 
71, no. 5, 1931, p. 475) and Hellmayr (Cat. Birds Ani jwoliayanmno: 
1937, p. 43) concluded that differences are due to individual variation. 
In a large series of specimens from Panama, as many have the tail 
longer than the wings as the reverse. 

The Yellow-billed Cacique inhabits dense thickets in forest openings 
and edge, second growth, abandoned fields, and, occasionally even man- 
groves, mostly at their edge or near islands of higher ground where 
there is some low vegetation. It is far more often noted by its calls than 
its appearance. In handling these birds I noticed that they lack the 
pungent odor of the other caciques and oropendolas. The male has a 
variety of pleasant whistling notes as well as the harsh, often explo- 
sive, calls that are given by both sexes; one that I noted was a wrenlike 
chatter, chruh-chruh-chruh. When seen, the birds often seem quite jay- 
like in movement, often resting with the tail hanging straight down. 
Rarely does one stay long in the open. So far as has been reported, 
they feed exclusively on insects. Ridgely (in litt.) has at least once 
seen a pair attending a small army-ant swarm. He has also seen them 
in flowering and fruiting trees, but it was not clear that they were feed- 
ing on anything but insects. One that I was able to watch closely at 
Chiva Chiva, Canal Zone, was climbing slowly about in heavy under- 
growth, keeping under cover. Several times I saw it pull masses of 
lodged dead leaves apart. Later it came out higher up on a slender 
dead branch that had some insect infestation. Repeatedly it thrust the 


FAMILY ICTERIDAE 355 


closed, chisel-pointed bill into the bark and then opened the two halves 
to pry off loose flakes. 

This species is found in pairs all through the year and is rarely seen 
in groups. Its nest is a cup, not a pendant bag like that of typical 
Cacicus. Hallinan (Auk, 1924, p. 320) found a nest at Gatun, Canal 
Zone, on April 24, 1909. It was “in a crotch of a limb in a sparsely- 
leaved shrub, in the jungle, about 8 feet from the ground. It was closely 
woven with heavy grass and light twigs and thickly lined with finer 
grass, with a cavity 3% inches in diameter and 2 inches deep. Two 
eggs, light blue, sparsely spotted with black, denser at the larger end.” 
Another nest, found at Summit, Canal Zone, on September 14, 1941 by 
Major General G. Ralph Meyer was of similar construction and con- 
tained two eggs that measured 27.2 < 18.0 and 27.7 X17.8 mm. In Costa 
Rica, Skutch (Pac. Coast Avif. no. 31, 1954, p. 284) found fledged 
young accompanying their parents in early March, indicating a breed- 
ing season beginning no later than January. A newly hatched nestling 
he found in May had pink skin with no down, tightly closed eyes, and 
red mouth lining. 

Eisenmann reports among other calls he has heard this species give 
in Panama, a clear, emphatic, whistled pur-weé-pew; a rather sweet 
wrreeeeoo; a harsher kweéyoo; a kwack-kwack-kwack, sometimes with 
a buzz in it like kzaak. 

Until recently this species was placed in a monotypic genus Ambly- 
cercus. 


SCAPHIDURA ORYZIVORA (Gmelin): Giant Cowbird, 
Vaquero Gigante 


FIGcuRE 29 


O.[riolus] oryzivorus Gmelin, 1788, Syst. Nat., 1(1), p. 386. (Cayenne.) 

Cassidix oryzivora violea Bangs, 1900, Proc. New Eng. Zool. Club, 2, June 30, 
p. 11. (La Concepcion, Santa Marta, Colombia.) 

Psomocolax oryzivorus impacificus Peters, 1929, Proc. Biol. Soc. Washington, 42, 
pulZoy Ceasa Nueva. Veracruz) Mexico.) 


Large; glossy black with violet sheen. 

Description.—Length, male 305-346 mm, female 270-315 mm. Adult 
male, entirely glossy black with violaceous sheen. Shows neck ruff in 
display. 

Adult female, much smaller, entirely black with less gloss and smaller 
ruffs than male. 

A male collected at Puerto Armuelles, Chiriqui, on March 1, 1966, 


356 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


had the iris light reddish brown; bill, tarsus, toes, and claws black. A 
female collected there that day was the same except for the iris, which 
was light mouse brown. At death, the reddish color of the iris changes 
within seconds to orange and then pale straw yellow (Olson, Proc. 
Wash. Biol. Soc., vol. 86, no. 34, 1973, p. 408). 

Measurements—Males (10 from Panama and Colombia), wing 
169.9-204.5 (191.1), tail 129.5-157.0 (145.4), culmen from base 35.9- 
39.9 (37.5), tarsus 41.7-48.0 (44.5) mm. 

Females (10 from Panama and Colombia), wing 145.5-167.0 
(155.2), tail 112.5-133.0 (119.2), culmen from base 31.3=33.3"(2208 
tarsus 37.2-39.9 (38.8) mm. 


Figure 29.—Giant Cowbird, Vaquero Gigante, Scaphidura oryzivora, male. 


Resident. Uncommon but widespread in lowlands of both slopes, 
except in the drier areas; it is absent from the eastern side of the 
Azuero Peninsula, southern Coclé, and most of western Province of 
Panama. In July 1975 and March 1981, Ridgely (in litt.) found this 
species more numerous in the Tuira lowlands of Darién than anywhere 
else in Panama. In July they were in flocks of well over 100 birds; in 
March, the breeding season, the birds were more dispersed, but equally 
abundant. This species parasitizes oropendolas and the colonial caci- 
ques, so during their nesting season is found in the same areas; at other 
times it is dispersed over more open country, generally feeding and dis- 
playing on the ground. 

The Giant Cowbird is found from southern Mexico to southeastern 
Brazil and northeastern Argentina. Peters (Proc. Biol. Soc. Wash., 


FAMILY ICTERIDAE 357 


vol. 42, 1929, p. 123) is responsible for dividing this species in Middle 
America, and he also recognized the race violea Bangs, which Hell- 
mayr (Cat. Birds Am., vol. 13, no. 10, 1937, p. 49) puts into oryzivora. 
The forms are supposedly separable on size, but Peter’s statistics are 
not at all convincing. The break between impacifica of southern Mexico 
and oryzivora is supposed to be in central Panama, but there are no 
detectable differences in size between birds from Chiriqui, Los Santos, 
the Canal Zone, Darién, or Colombia. A perfunctory examination of 
birds from north of Panama disclosed no significant size variation. 
Nor are color differences (more violaceous as against more bronzy 
gloss in the nominate race) apparent. Until there is good supporting 
evidence for recognizing geographic variation in this species, it should 
be regarded as monotypic. Parkes (Condor, 1954, p. 229) has shown 
why Scaphidura, not Psomocolax, is the correct prior name for this 
genus. 

The Giant Cowbird, formerly known in English as the Rice Grackle, 
is indeed sometimes found with cattle. It rides on their backs, where it 
picks off parasites, as well as taking insects flushed by the cattle as they 
move about. It also feeds independently in fields and pastures, on the 
exposed banks of rivers, and in trees. One I collected when it came 
to drink at a river in Los Santos had the throat filled with hulled rice 
eaten in a nearby field. Except for the brief season when these birds 
are depositing their eggs in the nests of oropendolas and colonial caci- 
ques, Giant Cowbirds are found in flocks in which females out-number 
males; often they feed with other icterids or with Groove-billed Anis 
(Crotophaga sulcirostris). Courtship occurs when the birds are in 
flocks: the males parade with the neck ruff distended and the bill bent 
down on the foreneck; sometimes when approaching a female, a male 
will bob up and down while in this posture. When skinning a male I 
found that the neck skin was loose so that it slipped readily over the 
head, and a sheath of dermal muscle was spread broadly over the dorsal 
and ventral sides. Presumably this is associated with the displaying of 
the ruff. 

Female Giant Cowbirds wait patiently around the colonies of oropen- 
dolas and caciques for the opportunity to deposit an egg in one of their 
nests. They are often driven away, but not so vigorously that it is im- 
possible for them to slip in and out of a nest. There are several de- 
scriptions of the eggs. Twelve taken from a colony of Cacicus cela in 
Suriname (Haverschmidt, Bull. Brit. Orn. Club, 87, 1967, pp. 136-137) 
were pale blue with a few small black spots and hair lines. Eight were 
elliptical, four oval. They measured 30.2-35.3X25.1-28.0 mm. The 


358 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


cacique eggs were white with similar black marks. Eggs taken from 
Psarocolius nests were longer, more elongated, with white ground color. 
An egg from Costa Rica was “spotless white, rough in texture and 
slightly glossed” and measured 36.1 26.0 mm (Crandall, Zoologica, 
vol. 1, no. 18, 1914, pp. 338-339). 

In Panama, N. G. Smith (Nature, vol. 219, 1968, pp. 690-694) 
found that eggs either mimicked in size and color those of their host or 
were “generalized” and non-mimetic. Response of the host species 
varied: at some colonies the hosts were more aggressive to visiting cow- 
birds and removed from the nest any eggs they detected as different 
from their own, while the entire population of other colonies was much 
more tolerant, neither driving away the cowbirds nor ejecting their 
eggs. Smith correlated tolerance with the absence of wasp or bee col- 
onies 1n the nest tree, since these insects somehow repel parasitic bot- 
flies (Philornis) that place their eggs or larvae on nestlings; the young 
cowbirds—themselves covered with down at hatching and less vulner- 
able to the botflies—preen their downless nestmates and eat any insect 
parasites they find. In the colonies lacking protective insects, produc- 
tivity of the host species was higher in nests with cowbirds than in 
those without. Smith suggested that the reason some female cowbirds 
continue to place their eggs in nests where they have a great chance of 
being rejected may be that those sufficiently mimetic also gain the pro- 
tection from other predators provided by the wasps or bees. 

The adult male Giant Cowbird is rarely seen at oropendola or cacique 
colonies, but fledglings often associate with young oropendolas and 
when begging for food may stimulate feeding by adult cowbirds if 
present (N.G. Smith in litt. to Eisenmann). 


MOLOTHRUS AENEUS AENEUS (Wagler): Bronzed Cowbird, 
Vaquero Bronceado 


Ficure 30 


Ps.[arocolius| aeneus Wagler, 1829, Isis von Oken, 22, heft 7, col. 758. (Laguna, 
Veracruz, Mexico.) 


Medium size; entirely black. 

Description.—Length, male 195-208 mm, female 175-184 mm. Adult 
male, entirely shiny black, iridescent purple on wings and tail. 

Adult female, entirely black, but with less iridescence than male; 
undersurface, especially at throat, often tinged dark brown. 

An immature male taken at Puerto Armuelles, Chiriqui, on February 


FAMILY ICTERIDAE 359 


8, 1966, had the iris light mouse brown; bill dark dusky neutral gray; 
tarsus, toes, and claws black. In adult males the iris is red. 
Measurements.—Males (10 from Panama and Costa Rica), wing 
111.0-120.6 (115.9), tail 68.8-82.0 (75.3), culmen from base 21.3-33.9 
(2245), tarsus 29.1-32:6'(31.0') mm. 
Females (10 from Panama), wing 95.5-106.0 (103.1), tail 65.4-72.1 
(69.5), culmen from base 18.8-20.8 (19.9), tarsus 27.4-28.6 (28.2) 


mm. 


A 


Oia. ee 
—- = oe 
Ps 


Figure 30.—Bronzed Cowbird, Vaquero Bronceado, Molothrus aeneus aeneus, 
female (left), male (right). 


Resident. Common in the lowlands of Bocas del Toro; local and less 
common on the Pacific slope from western Chiriqui to eastern Province 
of Panama, although never recorded in the Canal Zone. In recent years 
it has been recorded regularly and in substantial numbers in the lower 
highlands of Chiriqui; e.g., 45 at Volcan on March 20, 1979 (Ridgely 
in litt.). Ridgely (1976, p. 308) writes that ‘most central Panama re- 
ports are from the Tocumen/La Jagua area of eastern Panama Prov- 
ince. Several recent sightings from the Maria Chiquita/Rio Piedras 
area of eastern Colon Province (N.G. Smith et al.); also a flock of 8 
seen on eastern shore of Madden Lake on February 10, 1973 (Eisen- 
mann). Is perhaps becoming more numerous in central Panama, and 
spreading eastward.” This race is found north to south-central Texas. 
Other races occur in Arizona, Mexico, and Colombia. Parkes and Blake 
(Iieldiana: Zool., vol. 44, no. 22, 1965, pp. 207-216) have shown that 


360 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


this species lacks any characters important enough to justify separating 
it from other molothrine cowbirds in the genus Tangavius. They also 
argue for the inclusion of armenti Cabanis of northern Colombia as a 
race of aeneus. 

The Bronzed Cowbird is usually seen in open country where it feeds 
on the ground in fields and pastures, with or without cattle. It is often 
in flocks; one that I saw leaving a swampy roost at Tonosi, Los Santos, 
at sunrise, numbered 500 or so. On January 23, 1976, near El Rincon, 
Herrera, Ridgely noted over 600. If no conspecifics are available it 
may join flocks of other species—I once saw a cowbird at Chepo, Pan- 
ama, flying and feeding with a large band of Dickcissels (Spiza ameri- 
cana). A female Strauch (Bull. Brit. Orn. Club, 1977, p. 64) collected 
weighed 54.0 g. | 

The brilliant red of the iris of this species is probably due to perfu- 
sion with blood, since the eyes of 1 that I collected faded in half an 
hour to a deep reddish orange. 

In courtship the males spread the ruff on the sides of their neck, arch 
their wings, bend the head so the bill touches the breast, bring the tail 
forward, and fluff out their other body feathers so that their body re- 
sembles a sphere. Sometimes they fly around this way, and once, on 
March 3, 1960, at Buena Vista, near Concepcion, Chiriqui, I saw one 
all puffed out rise straight up a meter off the ground, where he hovered 
like a little helicopter in one spot for 10 or 15 seconds. To Eisenmann 
(in litt.), the display shows considerable resemblance to that of the 
Giant Cowbird. The male’s song is like that of the northern Brown- 
headed Cowbird (M. ater), but wheezier and throatier (Friedmann, 
1929, The Cowbirds, Springfield, Ill.: Charles C. Thomas, p. 325); the 
commonest call note of both sexes is a harsh chuck. When displaying, 
the male bounces up and down on the ground, meanwhile giving gut- 
tural yet bubbling notes, then a few high squeaky notes. 

The Bronzed Cowbird lays its eggs in the nests of small birds of 
many families. Friedmann (Smiths. Misc. Coll., vol. 149, no. 11, 1966, 
p. 7) says 56 species are known as hosts over the cowbird’s wide range; 
most build cup nests. Kiff (Wilson Bull., 1973, p. 73) lists 3 additional 
host species from Costa Rica. N. G. Smith (in litt. to Eisenmann) 
states that in the Almirante area of Bocas del Toro in late March 1965 
both this species and the Giant Cowbird were frequenting and appar- 
ently parasitizing oropendola colonies. Smith collected both kinds of 
cowbirds to prove the point. I have no other information on hosts in 
Panama. 


FAMILY ICTERIDAE 2061 


Friedmann (op. cit., 1929, p. 327) describes the eggs as “very light 
bluish green, bluish, or greenish, and without any markings whatso- 
ever’; their average measurements are 23 X18 mm. In southern Texas, 
the incubation period is 12 to 13 days and the young leave the nest at 11 
days, although they may be fed by their foster parents for another 2 
weeks. 


MOLOTHRUS BONARIENSIS CABANISII Cassin: Shiny Cowbird, 
Vaquero Brillante 


Molothrus Cabanisu Cassin, 1866, Proc. Acad. Nat. Sci. Philadelphia, 18, p. 22. 
(Santa Marta, New Grenada.) 


Medium size; male glossy violet-black; female drab grayish brown. 

Description.—Length, male 227-241 mm, female 196-216 mm. Adult 
male, body entirely very glossy violet-black; wings and tail more green- 
ish black. 

Adult female, entire upper surface grayish brown; wings and tail 
darker dusky brown; undersurface light brownish gray, paler on 
throat; back and breast more or less finely streaked with dusky. 

Immature, like female but more dusky above, the feathers edged 
with brownish buff; below more or less yellowish buff; males streaked 
below with dusky. 

A male collected at Fl Real, Darién, on February 17, 1964, had the 
iris dark brown; bill black; tarsus, toes, and claws black. 

Measurements.—Males (10 from Panama and Colombia), wing 
2379-15020) (112877), tail 97.0-107.2 (102.7), culmen trom base 19.5- 
21.8 (20.9), tarsus 26.6-29.2 (27.9) mm. 

Females (10 from Colombia), wing 98.6-111.5 (107.8), tail 78.5- 
89.1 (85.9), culmen from base 17.5-19.6 (18.6), tarsus 24.9-26.6 
(25.4) mm. 

Resident. Rare in eastern Darién and eastern San Blas. W. B. 
Richardson collected an immature male at Boca de Cupe on the Tuira 
River, Darién on April 3, 1915 (AMNH no. 136758). On February 
17, 1964, I shot an adult male from a treetop beside a cattle corral at 
Il Real, Darién, and a few days earlier I saw several along the Tuira 
River between Boca del Cupe and Yape. In March 1981, Ridgely (in 
litt.) saw up to 4 per day along the lower Tuira above FE] Real. From 
San Blas there is 1 sight report of a group of 7 or 8 birds, including 3 
or 4 males, seen by D. Sheets east of Puerto Obaldia in June 1965 
(Ridgely, 1976, p. 309). Ridgely (in litt.) saw 8 on the upper Rio 
Bayano, Province of Panama, between Aguas Clara and the Rio Diablo 


362 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


on April 24, 1976. This race is also found in adjacent Colombia, and 
other races are found to eastern Peru, southern Argentina, Chile, and 
southern Brazil. 

This species has been extending its range northward, having recently 
spread in the West Indies to the Lesser Antilles, Puerto Rico, and His- 
paniola. In early 1980, large numbers were seen around the Tocumen 
marsh and at the La Siesta Hotel, eastern Province of Panama, by 
many people (Ridgely in lit.). They were seen from at least January 
to March by observers previously familiar with both small cowbirds. 
The maximum count was approximately 200 on January 20, seen by 
P. Donahue, but usually the number seen was under 100. For a time 
the birds roosted at the hotel with Orchard Orioles. In the same period 
of 1981 neither cowbird was seen in this area. 

Friedmann (1929, The Cowbirds, Springfield, Ill.: Charles C. Thom- 
as, pp. 57-144) has studied this species in Argentina, where its vocaliza- 
tions are similar to those of M. ater of North America. Eisenmann (1m 
htt.) has heard a “pleasant tinkling song”’ from this species in Vene- 
zuela. It parasitizes a wide variety of small birds, but it has also oc- 
casionally been observed beginning the construction of a nest, although 
never finishing it. 


QUISCALUS MEXICANUS PERUVIANUS Swainson: Great-tailed 
Grackle, Changamé 


Quiscalus Peruvianus Swainson, 1838 (1837?), Anim. Menag,, p. 354. (Peru.) 


Very large; tail graduated; male entirely black with a glossy purple 
sheen; female dark brown above, lighter brown below. 

Description.—Length, male 389-414 mm, female 281-330 mm. Adult 
male, entirely black with a glossy purple sheen on body; wings and tail 
with slightly greenish sheen; tail long and keel shaped. 

Adult female, crown sepia, rest of upper surface, including wings 
and tail, very dark brown, slightly glossy; superciliary lighter brown; 
malar and auricular areas, and sides of neck sepia; chin dingy whitish, 
becoming buffy on throat; flanks, abdomen, and undertail coverts dark 
brown; rest of undersurface light brown, slightly lighter, more buffy 
on center of belly. 

A male taken at Chico, Panama, on March 16, 1949, had the iris yel- 
lowish white. The eye color of females varies from rather dull yellow 
to a light brownish yellow. In young birds the iris is dark. 

Measurements —Males (10 from Panama), wing 166.5-186.4 


FAMILY ICTERIDAE 363 


(175.1), tail 165.4-200.5 (184.2), culmen from base 40.3-45.7 (43.2), 
tarsus 43.3-48.8 (45.8) mm. | 

Females (10 from Panama), wing 139.9-150.2 (144.1), tail 132.5- 
155.9 (144.8), culmen from base 31.7-36.8 (34.6), tarsus 36.0-41.6 
(38.4) mm. 

Resident. On the Pacific slope this species is scarce in Chiriqui, but 
increasingly common as one moves east toward central Province of 
Panama and Darién. There are a few records from western Chiriqut: 
on March 9, 1966, I watched a male bathing in the Canal del Colorado 
at Puerto Armuelles, and on March 6, 1976, Ridgely (in litt.) saw 50 
in mangroves and on the seashore at I‘stero Rico. It is common on most 
of the islands off the Pacific Coast. On the Caribbean slope, it occurs 
quite commonly in the town of Changuinola, Bocas del Toro (April 
1980, Ridgely im litt.), and from just west of the Canal Zone eastward 
through San Blas and the islands off San Blas. It is most abundant in 
open residential areas and along the seashore. In Chiriqui and Darién, 
it seems as yet not to have spread far from its presumed ancestral habi- 
tat, mangroves and seashore areas. Beyond Panama this race is found 
on the Pacific Coast of Costa Rica and south to northwestern Peru and 
the Caribbean coast of Colombia and Venezuela. Other races are found 
north to southern Arizona and the Gulf Coast. 

Dr. Graham Fairchild of Panama City told me in 1960 that this 
species has increased greatly in abundance since he first came to Pan- 
ama 20 years earlier. With the increase in number, the birds are spread- 
ing back along the Canal. In 1960, I found a dozen at Gamboa; previ- 
ously I had never recorded any number beyond Fort Clayton and the 
Miraflores locks. Dr. Fairchild told me that the grackle roosts in the 
main plaza at Panama once became such a nuisance that the authori- 
ties finally cut down the trees. In contrast, Thayer and Bangs (Bull. 
Mus. Comp. Zool., vol. 46, 1906, p. 221) saw only 1, an adult female, 
in Panama City in 1904. They wrote that “the grackle is one of the 
birds relentlessly hunted for food by the natives, and is found, conse- 
quently, in very small numbers, and is exceedingly shy.” 

As might be expected of a bird that expands so successfully when not 
molested, the Great-tailed Grackle is opportunistic and aggressive. | 
have seen them feeding along the shore of rivers and oceans, taking 
fish (sometimes by diving a few inches), frogs, and crabs dead and 
alive, hawking for flying ants and termites, picking parasites off cattle, 
extracting insects from the ground, and eating berries. One that I 
watched eating ripe berries of a royal palm ( Roystonea sp.) seized them 


364 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


in the tip of its bill, pulled them off, and then manipulated them back 
into the mouth, where the berry was given a hard squeeze that crushed 
it. When it was rolled forward again, the pit was discarded with a 
scissorlike motion of the mandibles and the pericarp was swallowed. 
All this was done rapidly and expertly. Other foods I have not myself 
witnessed being taken are the eggs and young of other birds, and, not 
uncommonly, adult birds as large as a Groove-billed Ani. A female 
collected by Strauch (Bull. Brit. Orn. Club, 1977, p. 64) weighed 
1120) g: 

These are noisy birds with a wide repertoire of raucous calls: “both 
sexes have a harsh chack; male gives a long strident whistle, weeek, 
week, sometimes drawn out to weeeeek, and both sexes also give a rattl- 
ing trit-trit-trit’” (Ridgely, 1976, p. 309). It also has some more musi- 
cal calls. The males, which are only half or a third as numerous as the 
females, are aggressive about fighting among themselves and in court- 
ing females. Males court by fluffing out the body plumage, shaking the 
wings, and extending the bill and neck, while uttering a variety of calls. 
Once I saw a male following a female in flight suddenly hold the wings 
fully extended and flap them well below the line of his body for 8 or 10 
strokes. 

I have seen females gathering nest material on January 21, 1961, at 
Corozal in the Canal Zone and on February 16, 1944, at San José in 
the Archipiélago de las Perlas. Males that I collected on March 17, 
1946, at Jaqué, Darién, were only approaching breeding condition. On 
Taboga Island, Panama, I saw females carrying food to young in the 
nest on March 19, 1952, and on May 5, 1953, I saw grown young out 
of the nest at Balboa, Canal Zone. Dorothy M. Hobson (Year Book 
Indiana Audubon Soc., vol. 26, 1948, p. 24) saw young being fed in 
Panama City on July 20, 1947. Evidently the nesting season varies in 
date from locale to locale, and perhaps from year to year. 

Skutch (Pac. Coast Avif. no. 31, 1954, pp. 321-334) has studied this 
species in Guatemala, where it nests in colonies that are often placed in 
trees near water or in palms, as they are in Panama as well. However, 
they may at times nest nearer the ground. Major General G. Ralph 
Meyer found 3 nests on Taboguilla Island, Panama, on April 9, 1944, 
that were in low bushes not over 1.3 m from the ground. The nest is 
always a large, bulky cup made of grasses and other pliable plant fibers; 
as in most other colonial icterids, there is much attempted stealing of 
material from nearby nests, and I once saw a female simply take over 
a mockingbird’s nest under construction. A nest cup actually con- 


FAMILY ICTERIDAE 305 


structed by a grackle measured 10 to 13 cm in internal diameter and 6 
to 10 cm in depth. 

All the nests so far reported in Panama had a clutch of two eggs, 
although in Guatemala Skutch found three-egg clutches most common. 
The eggs have a “ground color .. . from bright blue to very pale bluish 
gray, on which are dots, blotches, and intricate scrawls of brown and 
black. The blue ground color of some eggs is locally washed with shades 
of brown or pale lilac. The diversities of pattern are so great that, if 
all the eggs in a populous colony were mixed together, each bird might 
conceivably be able to recognize her own by its distinctive markings. 
The measurements of 62 eggs .. . average 33.6 by 23.0 millimeters” 
(Skutch, op. cit., pp. 328-329). Incubation is 13 or 14 days, and, like 
all aspects of the nesting cycle except guarding the colony, is performed 
only by the female; both sexes, however, drive away hawks and other 
potential predators. The young hatch with pale salmon colored skin 
and sparse long gray down on the head, back, wings, and legs. They 
are fed insects and when 16 to 19 days old begin climbing outside the 
nest, although they can not yet fly. At this stage they resemble females, 
but with more grayish breasts, brown eyes, and faces and foreheads 
still unfeathered. When 20 to 23 days old they begin to fly, and con- 
tinue to follow their mothers for food for several days or weeks. In 
Panama the adults molt between September and November. 


ICTERUS SPURIUS SPURIUS (Linnaeus): Orchard Oriole, 
Parao de Huertos 


O.[riolus| spurius Linnaeus, 1766, Syst. Nat., ed. 12, 1, p. 162. (South Carolina.) 


Rather small; male with head, neck, upper back, wings, and tail black, 
rest of body rich reddish brown; female with upper surface grayish 
olive-green, undersurface bright greenish yellow. 

Description —Length 148-165 mm. Adult male, head, throat, upper 
breast, and upper back black; rest of body chestnut; lesser and middle 
upper wing coverts and all underwing coverts chestnut; greater wing 
coverts black, tipped whitish and brown; remiges black with outer webs 
narrowly edged whitish; tail black with all but central pair of rectrices 
tipped whitish. 

Adult female, upper surface grayish olive-green, brighter on crown 
and rump; undersurface dull greenish yellow; wings dusky; wing co- 
verts edged with whitish forming two wing bars; tail bright olive-green. 

Immature male, like female, but lores, front of face, and throat black; 


366 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


sometimes chestnut feathers on yellow undersurface; tail feathers 
black, edged green. 

Measurements.—Males (10 from North America, taken in April to 
June), wing 74.5-82.0 (78.7), tail 66.3-71.3 (69.2), culmen from base 
16.1-18.2 (16.9), tarsus 19.6-21.9 (20.8) mm. 

Females (10 from North America, taken in April to July), wing 
74.0-78.0 (75.9), tail 62.6-69.0 (66.4), culmen from base 15.5-17.7 
(16.6), tarsus 19.3-22.1 (20.5) mm. 

Migrant and winter visitor from the north, breeding in eastern 
United States and southeastern Canada. Common in the lowlands of 
both slopes (less so on the Pacific slope of the western provinces), 
where it has been recorded from late July until mid-April; occasionally 
it is found on migration in the highlands up to 1650 m (Ridgely, 1976, 
p. 309). It occurs in light woodlands and open areas with trees. W. W. 
Brown, Jr. (Thayer and Bangs, Bull. Mus. Comp. Zool., vol. 46, 1905, 
p. 157) collected an adult male on Saboga Island, in the Perlas Archi- 
pelago, on April 13, 1904, and Eisenmann observed an immature male 
(with black throat) on Isla Coiba on October 9, 1965, but it is not other- 
wise recorded from any of the islands off either coast. It winters from 
southern Mexico to northern Colombia and Venezuela. 

Most Orchard Orioles arrive in Panama in August and leave by mid- 
March. An indication of its abundance as a migrant in central Panama 
is Loftin’s report (in litt. to Wetmore) that he banded 68 on October 
18-19, 1963, 4.8 km east of Panama City. During the winter, it gathers 
in roosts that may be used in successive years; one in a tree in front of 
the Ancon Police Station in the Canal Zone held at least 100 birds when 
I saw it in February 1952 and December 1955. Another, near the old 
Tocumen airport, had 300-400 in December 1973-January 1974; these 
birds seem to have shifted to the La Siesta Hotel. Eisenmann notes 
that most of the Orchard Orioles he saw in central Panama during the 
northern winter were adult male, in chestnut plumage. 

When in Panama, the Orchard Oriole feeds to some extent on nectar, 
particularly that of the tree Erythrina fusca. 

Morton (Ann. Missouri Bot. Gard. 66, 1979, pp. 482-489) speculates 
that the identical color of the adult male Orchard Oriole’s body plum- 
age and the floral parts surrounding the nectar of Erythrina fusca in- 
dicates a coevolved relationship. He notes that adult male Orchard 
Orioles exclude females and immatures from these trees when they are 
in flower, which is at the time the orioles are most abundant in Panama. 
The Orchard Oriole is the only species that opens Erythrina flowers 
“correctly,” thereby getting pollen on its chest. The correctly opened 


FAMILY ICTERIDAE 367 


flower reveals a patch of chestnut-brown matching the adult male’s 
plumage and, Morton hypothesizes, “repels” the bird, so that 1t moves 
to unopened flowers in another Erythrina. 

At Almirante, Bocas del Toro, a male | watched feeding with North- 
ern Orioles (J. galbula) on the pink blossoms of a vine, pecked at the 
other orioles, and, when they did not respond, pulled at their rump 
feathers. The only time I saw this species feeding near the ground was 
in January 1961 at Corozal, Canal Zone, when I watched 4 or 5 search- 
ing the rank grasses that grew up one-third or so of the height of a 2.5-m 
chain-link fence. I have occasionally heard Orchard Orioles singing in 
Pepruary. Skutch (A Naturalist in Costa Rica, 1971, p. 104) has 
heard them sing in Costa Rica in late July and August and for 2 months 
before their departure in April; he noted that immature males still in 
the green plumage sang more than full adults. 


ICTERUS DOMINICENSIS PRAECOX Phillips and Dickerman: 
Black-cowled Oriole, Parao Encapuchado 


Icterus prosthemelas praecox Phillips and Dickerman, 1965, Wilson Bull., 77, p. 
298. (Almirante, Bocas del Toro province, western Panama.) 


Medium size; black, with bright yellow on lesser and middle wing 
coverts, bend of wing, lower back, and lower undersurface. 

Description.—Length 173-193 mm. Adult (sexes alike), entirely 
black, except for lesser and middle upper wing coverts, bend of wing 
and underwing coverts, lower back, and undersurface from belly 
through undertail coverts, which are bright yellow. 

Immature, as adult, but back yellowish olive-green. 

Measurements.—Males (5 from Bocas del Toro and Costa Rica), 
wing 88.1-91.0 (89.6), tail 89.3-91.7(90.3), culmen from base 20.1- 
20.9 (20.6), tarsus 20.5-23.1 (21.9) mm. 

Females (7 from Bocas del Toro and Costa Rica), wing 80.3-87.5 
(83.8), tail 82.2-90.6 (85.7), culmen from base 19.4-21.9 (20.4), tar- 
puis 21.1-23.1 (22.1) mm. 

Resident. Uncommon in lowlands of western Bocas del Toro and 
adjacent parts of Costa Rica on the Carribbean slope. Other races of 
the prosthemelas group are found on the Caribbean slope north through 
Central America into southeastern Mexico; and the dominicensis group 
is found on some islands of the Bahamas and Antilles. In Panama this 
oriole is usually found near river banks, in humid woodland borders, 
clearings with trees, and in banana plantations. Specimens have been 
collected at Changuinola, Almirante, Western River, and Chiriquicito 


308 BIRDS OF THE REPUBLIC OF PANAMA—PART 4 


(Peters, Bull. Mus. Comp. Zool., vol. 71, 1931, p. 344). Phillips and 
Dickerman described this race on the basis of the juvenal plumage, in 
which “the black of the throat patch [is] more extensive [than in J. d. 
prosthemelas|, extending on to the lower breast, and the interscapular 
region solid black, instead of yellow-green.” Postjuvenal plumages of 
most individuals are like those of J. d. prosthemelas, although Bangs 
(Proc. Biol. Soc. Wash., vol. 19, 1906, p. 111) notes that some indi- 
viduals from Panama to Honduras have the black extending beyond 
the breast and onto the flanks and rump. 

I have seen this species three times, at Almirante, during February 
and March 1958. A pair that I collected there on February 21 was in 
breeding condition. The only vocalization I heard was a metallic plink 
call note, but it also has a song that is “a sweet and rapid, but not very 
loud, series of whistles” (Ridgely, 1976, p. 310). In Costa Rica, 
Ridgely (1m litt.) has seen this species feeding in flowering Erythrina 
trees with migrant orioles. 

The nest of this oriole is a pouch suspended like a hammock from 
the underside of a broad leaf such as a banana or palm. The nest is 
attached by strands passed through holes made by the bird in the leaf. 
Two nests collected at Almirante on May 16 and June 9, 1962, were 
made of dark and light fibers and rootlets; the first nest was lined with 
brown plant down and was 80 mm deep. Neither nest contained eggs. 
Skutch (Pac. Coast Avif. no. 31, 1954, p. 266) found a nest in Hon- 
duras that had 3 young birds; after leaving their parents, the young 
form small flocks of their own age group. 

The Panama race has often been separated specifically with other 
Middle American subspecies from the West Indian J. dominicensis 
under the species name J. prosthemelas. 


ICTERUS AURICAPILLUS Cassin: Orange-crowned Oriole, 
Parao Coroninaranja 


Icterus auricapillus Cassin, 1848, Proc. Acad. Nat. Sci. Philadelphia, 3 (1847), 
p. 332. (Santa Marta, Colombia.) 


Medium size; back of head bright orange; forehead, throat, back, 
wings, and tail black; rest of body bright yellow. 

Description.—Length 179-190 mm. Adult (sexes alike), forehead, 
orbital region, throat, and breast black; crown, nape, and sides of head 
bright orange; upper back, tail, and wing, except for lesser and middle 
coverts, black; lesser and middle wing coverts, rump, and undersurface 
below breast orange-yellow; bend of wing and underwing coverts light 
orange-yellow. 


FAMILY ICTERIDAE 369 


Juvenile, crown and nape yellowish olive; back buffy olive; rump 
greenish olive; wings dusky; tail dusky, with feathers edged buffy 
olive; side of face, undersurface pale yellow, brighter greenish wash 
across breast; bend of wing and underwing coverts yellow. 

A male collected at El Real, Darién, on March 21, 1964, had the iris 
wood brown; bill black; tarsus and toes dark neutral gray; claws black. 

Measurements—Males (10 from Panama and Colombia), wing 
87.2-95.1 (91.0), tail 74.2-91.0 (83.3), culmen from base 19.2-21.0 

(20.1), tarsus 19.6-23.3 (21.4) mm. 
~ Females (10 from Panama and Colombia), wing 84